THE RUFFED GROUSE Ufi HISTORY ^ PROPAGATION ^ MANAGEMENT e At DIN It iUM^ tOftEftt W. OARROW FRANK C f DMINSTER WALTER r. CRiSSiY '#A M^ njfT^ > f A ^^'•"^ 3B3S^^E3E3EZ3EE3^^^^^^S0 3 Q Marine Biological Laboratory Library q 0 Woods Hole, Mass. [Q IB □ III a IB ■^^•^ Q ra [I] H) Presented by [0 m -'ew York ^tate Museum ji g Albany, N. i, ^ ffl Q H to [Q IB ISE3^^^E3^^^^^^^^£3I THE HUFFED GHOUSE Life History • I'rnpnjinlion • \\iiii,i} Wintering Yards AT THE Research Center Containing All-purpose Pens I'i.\( kd End TO End 459 The Wired-in Wintering Pen at the Research Center 160 The Wintering Pen is Divided into Sections in Low r.oMuis to Discourage the Birds FROM Chasing One Another 161 The Service Room is Constructed with Sliding Windows and Doors that Open Inwards to Facilitate Catching, Feeding and Watering the Bird*; Oitside Idl Even Hand-Raised Grouse Seldom Make Good Mothers in Captivity 474 "Still Air" Incubator with Turning Trays to Hold Incubating Eggs and Pedigree Trays into w 111(11 mi K<;(;s are Shifted Before Hatching 477 Chicks May be Idkntiiied by Attaching a Rand to Leg or Wing Before Transfkh- RING THEM IROM THE I'eDK.KEK 1I\T(;11ING TrAY TO THE BkoODEK ll'.l Shortly After Hatching the Chicks are Placed in v i!\TTi:in Hhixidkr 1}!3 The Colony Brooder House into which the Chicks vkk Mo\ kd khhm i hi: Umti.k^ Brooders when from Ten to Twelve Days t)Li) I5 \i.i.-PuRP0SE Pens, Scattered Over the Rearing Field. Make Idem, I mt> i\ WHICH TO Raise the Young Grouse Transferred from the Colony Broodik House I'^-' LIST OF ILLUSTRATIONS Page Birds May be Kept in the All-Purpose Pens in the Rearing Field Until Well INTO THE Fall 493 White or Orange-Colored Chicken Feathers, Wired Fast to the Tail Feathers OF THE Grouse Before Liberation, Aid in Checking on Sibseqient Movements 505 Liberated Grouse Soon Adapt Themselves to Wild Conditions Although Some Never Regain their Fear of Man so Char.acteristic of Their Wild Cousins 505 An Excellent Covert in which Mixed Woods are Edged with Overgrown Fields AND Broken. Here and There, by Small Slashings and Abandoned Lands. Such a Covert May Provide Large Crops of Both Grouse and Timber, if Skill- fully Maintained 588 Typical Abandoned Upland 598 An Ideal Edge Such as this is of More Value for Wildlife than as a Site on which to Plant Evergreens 600 New York State's Game Management Areas are Developed on the Multiple Use Principle 603 An Occasional Small Slashing Makes Extensive Woodlands More Attractive to Grouse 621 Winter Feeding Stations Attract Only an Occasional Bird 632 In Back Pastures, Many a "Grousy Nook" is Maintained by Moderate Grazing 641 In the Early Stages. Small Patches of Fall Feeding Groi nds May be Maintain- ed Rather Easily 643 When Allowed to Grow Up. Considerable Cutting is Necessary to Release Food Species from the Competition of Their Taller Neighbors. Without this Help. Apple and Sumach (indicated by arrows) Will Soon be Choked Out 643 Fortunate is the Covert Which Supports a Scattering of Apple Trees along its Edges 645 A Brushed-out Woods' Road is Attractive alike to Nesting Birds, Broods and Adults 645 Where Winter Shelter Is Scarce, a Spot Such as This May Well Be Thinned to Encourage the Evergreens Beneath 646 Slash Lanes, from 30 to 50 Feet Wide. Provide Excellent Summer Feeding Grounds for Broods and Fall Feeding Areas for Adults 61" An Ideal Small Slashing, Clean Cut. Irregular in Outline and with Evergreens Nearby 649 A Combination Such as This Offers a Challenging Opportunity to Produce Large Crops of Both Grouse and Timber 651 Heavily Pruned Plantations and Grouse Are Not Compatible 652 Where the Proper Seed Trees Are Present Nature May Often Be Depended Upon to Re-establish Satisfactory Woody Cover 656 Planting Spruce by the Center Hole Technique 658 Most Hardwood Trees and Shrubs Respond Best When Planted Directly Into Newly Plowed Furrows 659 White Pine, Interplanted in .\n Opening in Second-Growth Hardwoods. Here Provides Much Needed Winter Shelter for Grouse 66(1 Where the Crown Cover Is Fairly Open an Underplanting of Shade-Tolerant Spruce May Also Furnish Winter Shelter 661 I.IST OF II.I.VSTRATIONS Page An Excellent. Planted Shrub Border 663 This Shrub Border Resulted from Cutting a Band 20 Feet Wide Along the Edge OF the Woods 663 There Is a Sharp Contrast in Undergrowth Density and Diversity Between Heav- ily Pastured Woodlands and Those from Which Cattle Have Been Excluded 665 Grouse Survey Field Crew Lined up Ready to Enter a Covert on the Connecticut Hill Area 700 Typical Specimens of Grouse Egg Shell Rimains as Left by Various Nest Preda- tors 712 Remains of Adult Grouse Killed by an Accepitrine Hawk (probably Cooper's) Showing Head Still Attached to Carcass, Femur Picked Clean But Not Fractured, and Gizzard Uneaten 713 Winter Grouse Trapping Setup Showing (upper) Shelter with Bare Ground Be- neath and (lower) Wire Cage Trap With Captured Bird 716 Grouse Trapped at Nest and Marked with Colored Chicken Ff others Wired to Its Tail 717 Rear Portion of Body Cavity of a Bird of the Year (left) Showing Bursa as Com- pared with that of an Older Bird 718 90 LIST OF FIGURES Number Title Page 1 Range of the Ruffed Grouse 49 2 Distribution of Recent Records of Ruffed Grouse Over Its Former Range IN THE North-Central and Mid-Western States 51 3 Relative Abundance of the Ruffed Grouse Throughout Its Present Range 55 4 Structure of the Fresh and of the Developing Grouse Egg 74 5 Annual Chances in Shell Thickness of Grouse Eggs Collected from the Ithaca, New York. Region^1936-1940 75 6 Annual Changes in the Weight without the Yolk Sac of Newly Hatched Grouse Collected frow the Ithaca, New York, Region — 1936-1947 77 7 Grouse Development from Hatching to Adulthood 82 8 Progression of Wing Feather Development in Young Grouse 87 9 Relative Development at Weekly Intervals of Wing Feathers of Young Grouse 10 Average Seasonal Weights of 394 Adult and 108 Young, Wild Grouse from New York 92 11 Seasonal Weight-Health Relationship of Adult Grouse 96 12 Grouse Chick Calls ^^ 13 Parts of New York State Most Typical of the Three Major Grouse Habi- tat Regions and the Location of the Investigation's Study Areas in Each H ^ 14 Distribution of Adult Grouse Flushes on One Compartment of the Con- necticut Hii.L Study Area All Seasons— 1932-1940 171 15 Distribution of Adult Grouse Flushes on the Adirondack Study Area— All Seasons- 1932-1941 . I"-' 16 Food Groups Taken bv 1.093 Am lt Grouse \t \ vuious Seasons 211 17 Seasonal Variations in the Amount of Some Important Foods Consumkd in 1.093 Adult Grouse in New York 21i i.t Gkoi sk 489 42 Deviation from Mean of Sprim; 1'opulations on Connecticut Hii.i wd \ii- iRONDACK Areas— 193(1-1942 524 43 Grouse Brood Survival Recorded on Connectki i Hh i. vnd \dirondack Areas— 1930-1942 528 11 Rki.ationship of Number of Chicks Hatched to Number and Percent Sur- \ivi\(, at End of Summer on Connecticut Hill and Adirondack Areas 1930-1942 530 45 Relationship of Adult Sihvival to Fall Population Level on Connecticut Hill and Adirondack Areas— 1930-1942 533 46 Relationship of Fall Population Level to Number of Am : rs Lost on C.on- NECTUi T Hill and Adirondack Areas 1930-31 to 1941-42 534 17 Kelationshii* of F\i.l I'oin i.\tion Level to Proportion of Vni i.is l.nsi on Connecticut Hill AMI Vdirondack Areas — 1930-31 to 1')||.I2 5:U 18 Adult Mortality (ScpicniluT Id Sepienii>cri in Relation to the C.omposition OF THE FaI.I. I'oF'I I.ATION ON CONNECTICUT HiLL AND ADIRONDACK ArEAS— 1930-31 TO I')||-I2 536 UST OF FIGURES Number Title Pa^e 49 Relative Survival among Adult Grouse Adjusted to the Mean Size and Age Composition of the Fall Population on Connecticut Hill and Adiron- dack Areas— 1930-31 to 1941-42 537 50 Seasonal Distribution of Dead Grouse Found on Connecticut Hill Area— 1930-1942 537 51 Grouse Population Fluctuations Recorded from Year to Year during the Investigation on the Connecticut Hill and Adirondack Study Areas — 1930-1942 545 52 Average Life Equation of Increasing Grouse Population 547 53 Average Life Equation of Stable Grouse Population 548 54 Average Life Equation of Grouse Population Decreasing Primarily as a Result of High Over-winter Mortality 548 55 Average Life Equation of Grouse Population Decreasing Primarily as a Result of High Nest Mortality 549 56 Average Life Equation of Grouse Population Decreasing Primarily as a Result of High Brood Mortality 549 57 Relative Degrees of Fluctuation of Fall Grouse Population Densities on Connecticut Hill AND Adirondack Areas — 1930-1942 559 58 Trend of Fall Population Levels on the Connecticut Hill and Adirondack Areas with Smoothed Values Superimposed 565 59 Years of Principal Declines in the Abundance of Ruffed Grouse in Various Regions of Its North-eastern and North-central Range— 1900-1944 566 60 Relationship of Trend in the Average Density of the Fall Grouse Popula- tion on the Connecticut Hill Area to the Densities within the Vari- ous Compartments Comprising the Area 569 61 Trends of Fall Grouse Population Densities koh the \ \hii>i > Comi-vrt- MENTS of the CONNECTICUT HiLI. \rE\ 1930-1012 T ' 0 ()2 ()^E OF THE Many Possible Arrangements of ('over Types \\nHi\ a Covert WHICH Shoii.d Provide Faceptionm. Ci>NniTi()NS for Both Bird vnd HUNTKK 595 63 An Analysis ok the Productiveness of Some of the More Usual Comhinv- TioNs OF Cover that Make up Grouse Habitat in New York State 610 61 Theoretical Distribution or the Same Amount of Critical Grouse Cover Types to Illustrate the Importance of Proper Arrangement ()13 65 A Practical Illustration of the Effect of Cover Type Arrangement in De- termining THE Number of Grouse Habitats in a Covert 613 66 Simplest Design for Planting Open Land to Provide Grouse Cover 616 67 More Productive Design for Planting Open Land to Provide Good Grouse Cover f*'" 68 Grouse Habitat Development on Submarginal Land 619 69 Organization of a Covert for Both Grouse and Timber Production 625 70 Approximate Maximum Allowable Harvest by Hunting According to Dens- ity OF Fall Population (" ' 71 Topography of Connecticut Hill and Adirondack Study Areas 696 72 Relationship of Wooded and Overgrown Land to Open Land on Connecti- cut Hill ^y~ UST OF FIGURES Number Title Pa^e 73 Data Sheet Used to Hecoku Information Reuarijinc Aullt Grouse Flushed . 702 74 Data Sheet Used to Record Grouse Nest Information 703 75 Data Sheet Used to Record Grouse Brood Information 704 7ft Data Shket Used to Record Information REOARniNc Dead Grouse Found 705 77 Major Components of Grouse Skeleton 723 7{J Leg Musculature. Lateral View 729 79 Wing Musculature. Latekai. View or Deep Muscles 735 80 Dige.stive System 738 81 Longitudinal Section of L.4rge Intestine ., 739 82 Feather Tracts and Spaces of the Ruffed Grouse 712 83 Diurnal Trends in Rectal Temperature of Two Groups of .\dult Grouse, Group A of Which Was Exposed to an Additional Hour of Illumina- tion AT Night 7.52 84 Diurnal Trends in Rectal Temperature of Male and Female Adult Grouse 753 85 Effect of Excitement on Rectal Temperature .\nd Respir.\tion Rate of Adult Grouse 753 86 Trends in Rectal Temperature Following Wetting among Three Groups of Adult Grouse Exposed to Different Conditions of Air Temperature AND Humidity 754 87 Trends in Rectal Temperature Following Wetting among Diseased as Com- pared WITH Healthy Adult Grouse 755 oo Physiologic Reactions of Two Grouse to Repose 757 89 Holding Board Used in Measuring Physiologic Reactions Showing Head- Cover IN Place and Removed 7.iJ1 90 Trends in Rectal Temperature among Two Group.-; of Fastini. Am i.t Grouse Held at Different Air Temperatures 702 91 Trends in Rectm. Temperature among Male and Femm.i: \\n ut (wjoise Fvst- iN(. withoit Drinking Water Td.'^ 02 Trends of 21-Hour Ciian(;es in Body Weight of Adi i.t (Jrouse during Pro- longed Fasting with and without Water 7()(i 93 Trends OF Percentage Loss IN Body Weight ok Adi i.t Grouse di rin(, Pkd- i.on(;ed Fastin(; with and withoit Water 7o7 91 Relationship between Body Weight Loss and Water Consumption of Vdui.t Groise DURiNi; Prolonged Fasting at Two Air Temper.\tures 770 95 Comparative Trends in Body Weight among Two Groups of Adult Grouse DURING Period of Fasting and Following Subsequent Ingestion of Food 772 121 Tm'i: of Cover Preferred by Nesting Ruffed Grouse — Entire St.\te — 1930- 19;K) "8.-! 122 Location of {{iffed (Arouse Nests as Influenced u\ Density of Slrhoumuni. Undergrowth in Various Types of Cover— Entire State — 1930-1936 7!:i. 123 Site Preference of Nesting Ruffed Groise — Entire St.^te — 1930-1936 785 124 Location of Ruffed Grouse Nests as Influenced by Type of Cover in Rel.\- TioN TO Degree of Slope — Entire State 1930-1936 Tl'.d 125 Location of Ruffed Grouse Nests as Influenced by Type of Comk in 1!el\- TioN TO Aspect of Slope — Entire State — 1930-1936 787 * The foMuwind finuro niimluT* iirc iiiaili; tit mnforiii »illl llir table number* with which they are aMOciatrd. LIST OF FIGURES Number Tide Page 126 Location of Ruffed Grouse Nests as Influenced by Distance from Nearest Conifers— Entire State— 1930-1936 789 127 Location of Ruffed Grouse Nests in Relation to Distance from an Open- ing—Entire State— 1930-1936 "90 128 Location of Ruffed Grouse Nests as Influenced by Type of Cover in Rela- tion TO Distance from the Nearest Slashing — Entire State — 1930-1936 791 129 Location of Ruffed Grouse Nests as Influenced by Type of Opening in Re- lation to Distance from an Opening— Entire State — 1930-1936 793 130 Fate of Grouse Nests According to the Type of Crown Cover in Which Found as Influenced by Region— Entire State— 1930-1936 794 131 Fate of Ruffed Grouse Nests as Influenced by the Undergrowth Density IN Various Types of Crown Cover— Entire State— 1930-1936 796 132 Fate of Ruffed Grouse Nests as Influenced in the Density of Cover at the Nest Site— Entire State— 1930-1936 797 133 Fate of Ruffed Grouse Nests as Influenced by their Distance from a Road, Trail or Other Opening in the Various Types of Crown Cover— Entire State— 1930-1936 '^98 134 Fate of Ruffed Grouse Nests as Influenced by the Type of Opening in Re- lation TO Distance from an Opening — Entire State — 1930-1936 800 1 35 Type of Cover Used by Grouse Broods at Various Ages — Entire State— 1930-1936 «0l 1 36 Density of Crown Cover Used by Grouse Broods at Various Ages — Entire State— 1930-1936 802 137 Type of Cover Used by Grouse Broods as Related to Time of Day — Entire State— 1930-1936 803 138 Type of Cover Used by Grouse Broods under Various Temperature Condi- tions— Entire State — 1930-1')36 804 139 Type of Cover Used by Grouse Huodds under Various Wind Conditions— En- tire State— 1930-1936 805 110 Type of Cover Used by Grouse Broods i nder Various Atmospheric Condi- tions—Entire State— 1930-1936 806 1 11 Type of Cover Used by Grouse Broods under Different Ground Conditions —Entire State— 1930-1936 807 1 12 Type of Undergrowth Used by Grouse Broods at Various Ages — Entire State- 1930-1936 808 143 Density of Undergrowth Used by Grouse Broods at Various Ages — Entire State- 1930-1936 "09 144 Type of Undergrowth Used by Grouse Broods under Various Temperature Conditions — Entire State — 1930-1936 810 145 Type of Undergrowth Used by Grouse Broods under Various Atmospheric Conditions — Entire State — 1930-19-36 811 146 Type of Undergrowth Used by Grouse Broods under Various Wind Condi- tions— Entire State — 1930-1936 812 147 Degree of Slope Used by Grouse Broods at Various Ages — Entire State— 1930-1936 813 LIST OF FIGURES Number Title t^cfi'' 148 Aspect of Slope Used by Grouse Broods as Related to Time of Day— En- tire State— 1930-1936 "!•■ 149 Aspect of Slope Used by Grouse Broods under Various Temperature Condi- tions—Entire State— 1930-1936 !!••■' 150 Aspect of Slope Used by Grouse Broods under Various Wind Conditions- Entire State— 1930-1936 816 151 Aspect of Slope Used by Grouse Broods under Various Atmospheric Condi- tions—Entire State— 1930-1936 817 152 Locations Used by Grouse Broods as Related to Interspersion of Crown Cover Types— Entire State— 1930-1936 818 153 Type of Cover Used by Adult Grouse by Seasons — Connecticut Hill Area —1930-1936 819 154 Type of Cover Used by Adult Grouse by Months— Connecticut Hill Area —1930-1936 822 155 Type of Cover Used by Adult Grouse in Winter as Related to Time of Day —Connecticut Hill Area— 1930-1936 823 156 Type of Cover Used by Adult Grouse in Spring as Related to Time of Day —Connecticut Hill Area— 1930-1936 825 157 Type of Cover Used by Adult Grouse in Summer as Related to Time of Day —Connecticut Hill Area— 1930-1936 827 158 Type of Cover Used by Adult Grouse in Fall as Related to Time of Day — Connecticut Hill Area— 1930-1936 829 159 Type of Cover Used by Adult Grouse in October and November as Related TO Time of Day— Entire State— 1930-1936 830 160 Type of Cover Used by Adult Grouse under Various Temperature Condi- tions— Connecticut Hill Area —1930-1936 831 161 Type of Cover Used by Adult Grouse under Various Wind Conditions — Con- necticut Hill Area— 1930-1936 ' 832 162 Type of Cover Used by Adult Grouse under Various Atmospheric Condi- tions— Connecticut Hill Area — 1930-1936 833 163 Type of Cover Used by Adult Grouse under Various Ground Conditions- Connecticut Hill Area— 1930-1936 834 164 Seasonal Preference of Adult Grouse for Ground or Tree — Four Areas — 1930-1936 836 165 I'reference of Adult Grouse for Ground or Tree in Various Types of Cover —Four Areas— 1930-1936 837 166 Aspect of Slope Used by Adult Grouse as Related to Time of Day— Entire State— 1930-1936 83}", 167 Aspect of Slope Used by Adult Grouse under Various Temperature Condi- tions—Entire State— 1930-1936 839 Id!! Influence of Cover on the Flushing Distance of Adult Grouse from tiik Observer— Four Areas— 1930-1936 !''"' 170 Influence of Sex on the Distance between Adult Grouse Flush and Re- flush— Entire State -1930-1936 843 171 Likelihood of Flushed Adult Grouse Alighting on Ground or in a Tree According to the Coveu Type Four Areas— 1930-1936 814 LIST OF TABLES Number Title Page 1 A Chronology of Local ok General Causes of Grouse Scarcity as Listed by Two OR More Authors 14 2 Records and Results of Grouse Liberations Known to the Lnvestigation IH 3 Known Attempts to Rear Ruffed Grouse in Captivity from Eggs Collected in the Wild— 1879-1942 22 4 Known Attempts to Rear Ruffed Grouse in Captivity from Eggs Secured FROM Hand-Raised Birds — 1875-1942 24 5 Recent Ruffed Grouse Investigations 30 6 Comparison of Visual Sex Differences in Ruffed Grouse 45 7 A Generalized Breeding Behavior Pattern Covering Adult Male Ruffed Grouse in Captivity Compiled from a Study of 78 Individual Patterns 69 Ji Physico-Chemical Properties of Grouse Eggs Collected from the Ithaca, New York. Region— 1936-1940 74 9 Chemical Composition of 189 Newly Hatched Grouse Chk k> Collected FROM the Ithaca, New York, Region— 1936-1940 76 10 Average Monthly Weights of 394 Adult and 108 Young Grouse Collect- ed in New York State— 1931-1941 '^1 11 Average Weekly Weights, in Grams, of 8 Male and 7 Female Hand-Reared Grouse Chicks 94 12 Length of Central Tail Feathers in Adult Grouse from New York State 98 13 Explanation of the Cover Types and Symbols Used in the Ruffed Grouse Investigation 120 14 Type of Cover Used by Grouse Broods Connecticit Hill Area— 1930-1936 140 15 Types of Cover Used by Adult Grouse— Connecticut Hill Area— 1930-1936 152 16 Relationship of the Frequency with which Edges Occur to the Average Level OF Spring Populations ON 12 Sections of the Connecticut Hill Area— 1930-1942 . 172 1 7 Relationship of the Number of Cover Types within 100 Feet of Broods Contacted to the Probability of Contact as Determined by Random Sampling — Connecticut Hill Area — 1930-1942 172 18 Relation of Grouse Flushes to Edges in Planted Coniferous Plantation and IN Natural Woodlands — Luther Preserve — 1933-1934 174 19 Major Studies of the Food Habits of Ruffed Grouse 183 20 Seasonal and Regional Distribution of the 1.093 Adult Grouse and 540 Chicks. Collected and Analyzed for Food Habits Studies — Entire State— 1931-1941 184 21 Average Volumes, in Cubic Centimeters, of the Crop and Gizzard Contents of 1,093 Adult Grouse, by Season and Relation — New York 186 22 Seasonal Variation in the Number of Food Items Per Individual Crop and Gizzard Taken by Grouse in New York 191 23 Volumetric Percentage of Gravel in the Gizzard of 1.093 Adult and 540 Chick Grouse, for Various Seasons of the Year and Regions of the State 193 24 Properties of Some Poisonous Plants Eaten by Grouse 195 IJST <)l TABLES Xuinber Title P Sources of Plant and Animal Foods that Bulked Largest in the Diet ok 1,093 Adult Grouse in New York — Summary of All Seasons — 1931-1941 19ii 26 Quantities OF Beechnuts Consumed BY 1.093 Adult Grouse in New York — 1931-1941 202 27 The Ten Groups of Plant Foods Most Commonly Eaten by Grouse in New York during Each Season, Showing the Volumetric Percentages, Im- portant Species and Parts Eaten 215 28 Regional Variation in the Ten Most Important Plant Foods of Adult Grouse for Each Season — New York 218 29 Yearly Variations in Percentage Bulk of Important Food Plants of Adult Grouse in New York in Comparison with the 11 -year Average 219 30 A Comparison of the Volumetric Percentages of Fall Foods Chosen by Adult Grouse in New York with that Reported from Other Parts of Its Range 220 31 Monthly Variations in the Important Plant and Animal Foods of 540 New York Grouse Chicks 224 32 Characteristics of Some Important Food Plants Commonly Utilized by Grouse in New York 227 33 Distance between Successive Nests of Individual Marked Female Grouse 259 34 Relation of Drumming Logs to Nests — Connecticut Hill — 1930-1942 266 35 Grouse Nest Mortality Recorded in New York State and Proportion Resulting from Predation — 1930-1942 311 36 Grouse Nest Mortality Recorded in Adirondack Region — 1931-1942 312 37 Brood Mortality Recorded on Connecticut Hill and Adirondack Study Areas— 1930-1942 315 38 Adult Mortality Recorded on Connecticut Hill and Adirondack Study Areas— 1930-1941 318 39 Incidence of Grouse Remains in Fox Droppings — Connecticut Hill Area — 1930-1941 322 10 Proportion of Broken-lp Grouse Nests Attributed to Various Predators IN Different Regions of New York — 1929-1942 334 11 Predators Considered Responsible for Dead Grouse Chicks Found on Con- necticut Hill and Adirondack Study Areas — 1930-1942 335 42 Predators Considered Responsible for Dead Adi lt Grouse Examined hv the Investigation — 1930-1942 337 43 Proportion of Different Types of Food in Predator Stomachs Examined BY THE Investigation — 1930-1942 339 44 Proportion of Diffp:rent Types of Food in Fox Droppings Examined by the Investigation 1930-1942 340 45 Proportion of Different Types of Food in Pellets of thk Ghkvt Mornkd Owl Examined by the Investk.vtion 1930-1942 312 16 Comparison of Nest Mortai.it\ on \rea Si h.iected to i*KKn\ToK Control and on Check Area — Connecticut Hill— 1931-1935 346 47 Comparison of Brood Mortality on Area Subjected to I'redator Control and on Check Area — Connecticut Hill — 1931-1935 347 USr OF TABLES » Number Title Page 48 Comparison of Adllt Mortality on Area Subjected to Predator Control AND ON Check Area — Connecticut Hill — 1931-1935 348 49 Relationship of Density of Grouse Breeders to that of Chicks at Hatch- ing AND to that of ToTAL POPULATION IN FaLL ON CONTROLLED AND ChECK Areas during Predator Control Experiments on Connecticut Hill — 1931-1935 349 50 Average Number of Eggs in First Nests — Entire State — 1931-1941 361 51 Greatest Variation in Average Annual Clutch Size (in eggs I 362 52 Average Number of Eggs in Renests — Entire State — 1931-1941 364 53 Infertility Rates in First Nests and Renests — Entire State — 1931-1941 365 54 Embryo Mortality in First Nests and in Renests — Entire State — 1931-1941 366 55 Reported Kill of Grouse and Average Seasonal Bag per Licensed Hunter Reporting in New York— 1923-1939 372 56 Grouse Killed by Hunters on Hunting Check Areas — 1930-1931 373 57 Analysis of Hunting Season Losses of Grouse in New York — 1930, 1931 and 1936 376 58 Crippling Loss on Population Control Experiment — Connecticut Hill 378 59 Success Ratio in Grouse Hunting — Tompkins County and Public Hunting Ground Hunter-Check Areas 378 60 Reported Take of Certain Predatory Fur Bearers in New York — 1926-1939 382 61 Ruffed Grouse Data — Pharsalia Refuge and Chenango Public Hunting Ground Surveys — February 1935-1937 391 62 The Number of Adult Grouse and of Grouse Chicks Collected kh; I'vtmo- LOGicAL Examination by Years AND Regions 403 63 The Animal Parasites of the Ruffed Grouse and their Rkpoiitki) Distiuhi - tion 109 64 Relative Occurrence of Parasites in Young and Adult Grouse in New York —1931-1941 410 65 Average and Maximum Numbers of Parasites Encountered in Grouse in New York— 1931-1941 411 66 Relative Occurrence of Pathological Conditions in Adult and Young Grouse in New York— 1931-1941 411 67 Incidence of the Common Parasites of Grouse in New York by Ace Classes —1931-1941 112 68 Penning and Productivity of Bobwhite (^uail Compared with Ruffed Grouse in Captivity 458 69 Average Weights in Grams of Hand-Reared Grouse at the Research Center DURING Certain Periods 463 70 Suggested Conditions for the Incubation of Grouse Eggs 478 71 Indices of Production 498 72 Summer Sex Ratios of Adult Grouse as Indicated by Population Estimates —Connecticut Hill Area— August 31, 1930-1942 514 73 Summer Sex Ratios of Adult Grouse as Indicated by Population Estimates —Adirondack Area— August 31, 1932-1942 515 LIST or TABLES Number Title ' Page 71 Sex Ratios of Grouse Collected during Mearly Complete Elimination of Population on Tract Adjacent to Connecticut Hill — October-March, 1934-1936 515 75 Sex Ratios of Immature Grouse — Statewide Collections — August 1 Through December 31. 1938-1941 510 76 Number of Females per lUO Acres in Spring Grouse Population on Connec- ticut Hill and Adirondack Areas— 1930-1942 51.5 77 Number of Grouse Broods per 100 Acres on Connecticut Hill and Adiron- dack Areas— 1930-1942 518 78 Number of Adult Grouse per 100 Acres on Connecticut Hill and Adiron- dack Areas— 1930-1912 519 79 Relative Security of Fall Grouse Populations on Connecticut Hill and Adirondack Areas— 1930-1941 523 80 Proportion of Broken-up Grouse Nests Observed during the Investigation Attributed to Various Causes — 1930-1942 527 81 Brood Survival Recorded on Connecticut Hill and Adirondack Study Areas —1930-1942 '527 82 Relationship of Density of Grouse Chicks at Hatching Time to Survival DURING Brood Period on Connecticut Hill and Adirondack Study Areas —1930-1942 529 83 Relationship of Density of Breeding Populations to Survival during the Brood Period on Connecticut Hill and Adirondack Study Areas — 1930- 1942 531 84 Adult Survival Recorded on Connecticut Hill and Adirondack Study Areas —1930-1941 531 '-^ 105 Average Standard Quiet Heart Rate of Inactive Adult Grouse 100 Hours AFTER Withdrawal of Food '•'^'-* 106 Calculation of Amount of Dependence between Physiological Reactions OF Feeding Adult Grouse 760 107 Calculation of Amount of Dependence between Physiological Reactions of Fasting Adult Grouse "60 100 Effect of Starvation with Water Avam.able upon the Rectal Tempera- ture of Adult Grouse '61 1(19 Effect of Starvation upon Respiration Rate of Adult Grouse 7()2 110 Effect of Starvation without Water upon the Rectal Temperature and Respiration Rate of Adult Grouse "63 1 11 Comparison of Body Temperature of Feeding Grouse with That of Grouse AFTER Fasting 84 Hours Both with and without Water 764 112 Loss in Weight among Adult Grouse during Starvation under Different Conditions '6o 113 Rate of Digestion of Adult Grouse Feeding Uninterruptedly at Different Air Temperatures '67 114 Rate of Digestion of Adult Grouse after Fasting at Different Air Temper- atures '68 115 Relationship between Environmental Temperature and Digestive Activity 768 116 Daily Caloric Intake in Relation to Weight Change of Adult Grouse at Two Air Temperatures 771 1 17 Daily Caloric Intake in Relation to Weight Change of Adult Grouse after Prolonged Fasting ''1 118 Calculated Daily Caloric Intake of Food Needed by Adult Grouse to Main- tain Weight and Vigor '"■^ 119 Numbers of Insects and Allied Groups, by Orders, Collected from 3-foot AND 10-FOOT Plots on the Connecticut Hill Area between June 2 — LIST OF JAliLES Number Title /'age July 1, 1936 AND June 1— July I. 1937; in Vicinity of tiik Adirondack Area between June 9— July 11. 1936 'i'TS 120 The Average ^UMBER of Insects and Allies Collected from 3-foot and 10- foot Plots in Different Types of Cover on the Connecticut Hill Area BETWEEN June 2— July 1, 1936; in Vicinity of the Adirondack Area between June 9— July 14, 1936 ' '6 121 Type of Cover Preferred by Nesting Ruffed Grouse — Entire State— 1930- 1936 '^3 122 Location of Ruffed Grouse Nests as Influenced by Density of Surrounding Undergrowth in Various Types of Cover— Entire State— 1930-1936 784 123 Site Preference OF Nesting Ruffed Grouse— Entire State— 1930-1936 785 121. Location of Ruffed Grouse Nests as Influenced by Type of Cover in Rela- tion TO Degree of Slope— Entire State— 1930-1936 780 125 Location of Ruffed Grouse Nests as Influenced by Tyte of Cover in Rela- tion TO Aspect of Slope— Entire State— 1930-1936 '8, 126 Location of Ruffed Grouse Nests as Influenced by Distance from Nearest Conifers— Entire State— 1930-1936 "89 127 Location of Ruffed Grouse Nests in Relation to Distance from an Open- ing—Entire State— 1930-1936 "'^" 128 Location of Ruffed Grouse Nests as Influenced by Type of Cover in Rela- tion TO Distance from the Nearest Slashing — Entire State — 1930-1936 791 129 Location of Ruffed Grouse Nests as Influenced by Type of Opening in Re- lation TO Distance from an Opening — Entire State— 1930-1936 792 130 Fate of Grouse Nests According to the Type of Crown Cover in Which Found as Influenced by Region— Entire State— 1930-1936 794 131 Fate of Ruffed Grouse Nests as Influenced by the Undergrowth Density in Various Types of Crown Cover— Entire State— 1930-1936 796 132 Fate of Ruffed Grouse Nests as Influenced by the Density of Cover at the Nest Site— Entire State— 1930-1936 "''" 133 Fate of Ruffed Grouse Nests as Influenced by their Distance from a Road. Trail or Other Opemn(. in the Various Types of Crown Cover -Entire State— 1930-1936 '^^^ 134 Fate of Ruffed Grouse Nests as Influenced by the Type of Opening in Rela- tion to Distance from an Opening— Entire State— 1930-1936 800 135 Type of Cover Used by Grouse Broods at Various Ages — Entire State — 1930-1936 ^^^ 136 Density of Crown Cover Used by Grouse Broods at Various Aces— Entire State— 1930-1936 ^^^2 137 Type of Cover Used by Grouse Broods as Related to Time of Day— Entire State— 1930-1936 ^^'^ 138 Type of Cover Used by Grouse Broods under Various Temperature Condi- tions—Entire State— 1930-1936 804 139 Type of Cover U.sed by Grouse Broods under \.\rious Wind Conditions— En- tire State— 1930-1936 ^•'■'' 140 Type of Cover Used by Grouse Broods under Various Atmospheric Condi- tions—Entire State- 1930-1936 8<'<> LIST OF TABLES Number Title Page 141 Type of Cover Used by Grouse Broods under Different Ground Conditions —Entire State— 1930-1936 807 142 Type of Undergrowth Used by Grouse Broods at Various Aces — Entire State— 1930-1936 808 143 Density of Undergrowth Used by Grouse Broods at Various Aces — Entire State— 1930-1936 809 144 Type of Undergrowth Used by Grouse Broods under Various Temperature Conditions — Entire State — 1930-1936 810 145 Type of Undergrowth Used by Grouse Broods under Various Atmospheric Conditions — Entire State — 1930-1936 811 146 Type of Undergrowth Used by Grouse Broods under Various Wind Condi- tions—Entire State— 1930-1936 812 147 Degree of Slope Used' by Grouse Broods at Various Aces — Entire State — 1930-1936 813 148 Aspect of Slope Used by Grouse Broods as Related to Time of Day — En- tire State— 1930-1936 814 149 Aspect of Slope Used by Grouse Broods under \ arious Te.mpekatlre Condi- tions— Entire State — 1930-1936 81o 150 Aspect of Slope Used by Grouse Broods under Various Wind Conditions — Entire State— 1930-1936 816 151 Aspect of Slope Used by Grouse Broods under Various Atmospheric Con- ditions— P^NTiRE State — 1930-1936 817 152 Locations Used by Grouse Broods as Related to Interspersion of Crown Cover Types ^Entire State— 1930-1936 818 153 Type of Cover Used by Adult Grouse by Seasons — Connkctk i t Hill Arex —1930-1936 819 154 Type of (]over Used by Adult Grouse by Months — Connecticut Hill Area —1930-1936 821 155 Type of Cover Used by Adult Grouse in Winter as Related to Time of Day —Connecticut Hill Area -1930-1936 823 156 Type of Cover Used by Adult Grouse in Spring as Related to Time ok Day —Connecticut Hill Area— 1930-1936 825 157 Type of Cover Used by Adult Grolse in Summer as Related to Time of Day— Connecticut Hill Area— 1930-1936 827 158 Type of Cover Used by Adult Grouse in Fall as Related to Time of Day — Connecticut Hill Area —1930-1936 829 159 Type of Cover Used by Adult Grouse in October and November as Relat- ed to Time of Day — Entire State — 1930-1936 830 1(>0 Type of Cover Used by Adult Grouse under Various Temperature Condi- tions— Connecticut Hill Area — 1930-1936 831 161 Type of Cover Used by Adult Grouse under Various \^'imi Conditions — Connecticut Hill Area— 1930-1936 832 1()2 Type of Cover Used by Adult Grouse under Various Atmosphekic Condi- tions— Connecticut Hill Area — 1930-1936 833 163 Type of Cover Used by Adult Grouse under Various Ground Conditions — Connecticut Hill Area — 1930-1936 831 LIST Ol- TABU:S Number Title Page 164 Seasonal Preference ok Adilt Grouse for Ground or Tree — Four Areas — 1930-1936 836 165 I'khference of Adult Grouse for Ground or Tree in Various Types of Cover —Four Areas— 1930-1936 837 166 Aspect of Slope Used by Adult Grouse as Related to Time of Day — Entire State— 1930-1936 838 167 Aspect of Slope Used by Adult Grouse under Various Temperature Condi- tions-Entire State— 1930-1936 839 168 Influence of Cover on the Flushing Distance of Adult Grouse from the Observer— Four Areas— 1930-1936 810 169 Type of Cover Used by Adult Grouse before and after Being Flushed — Four Areas— 1930-1936 842 170 Influence of Sex on the Distance between Adult Grouse Flush and Re- flush — Entire State — 1930-1936 'e\k on the Connecticut Hu.i, \mi Vdikdn- DACK Study Areas 1930-1942 !»!- 185 Seasonal Population Estimates for the Various Compartments of hie Con- necticut Hill Study Area 1930-1912 882 18() Pkopa(;ation Information on Trees & Shrubs Kecummemied i ok Plantations 883 PREFACE 'To know the grouse is to love it And loving it. to wish it uell." In a scientific report such as this, there is little room for personalities or individual feel- ings. Only in the preface can one give expression to thoughts quite unscientific but unde- niably human. When those we love are in trouble we do our best to help. So it is wilii tiic ruffed grouse. Two and a quarter centuries have passed since the first law protecting grouse was enacted in the Colony of New York. Since then a million men. gun in hand, heart in the highlands and the swish of leaves in their faces, have roamed woods roads and forest glades, hunting partridge and finding zest in life. That all who know the grouse come to have an affection for this unpredictable thunder- bolt of the uplands, is axiomatic. Its rolling drum and roaring flush add a personality to the woods which nothing can replace. Its uncanny skill in foiling the sportsman has endeared it to all who prize a worthy opponent. The partridge asks no favors. It has never been on relief, as with pheasants and quail in a northern winter. Yet. that it. loo. has trouble was realized years ago. The mysterious way in which, from abundance it suddenlv becomes scarce, then gradually grows plentiful again, has long caused genuine concern. In fact, these violent periodic fluctuations in grouse pop- ulation have made many fear for the future of this remarkable bird. More than a few endeavored to do something about it: Hodge. Torrey. Stoddart. Foriiush. Walcott, Burnhani. Allen and a host of others: thus the honor roll runs. As far back as 1907 the New York State Conservation Department realized the need for an investigation of these so-called epidemics. Accordingly, it employed E. Seymour Wood- ruff to send out questionnaires to obtain the opinion of sportsmen and game protectors throughout the State as to the cause. A. M. Stoddart. Rod and Gun editor of the New York Sun, made a similar study for the Department covering the 1917 scarcity. For over ten years it cooperated with Professor A. A. Allen of Cornell University by sending him, through its game protectors, the viscera of grouse from all over the State to facilitate his study of grouse diseases and by encouraging his pioneer researches into the artificial raising of this species. Though plent) had always followed scarcity, by 1928 grouse were at such a low ebb in New York that a closed season was invoked and retained through 1929. In August of that year, a group of earnest, worried sportsmen, led by George Lawyer of \^'atertown. met with Conservation Commissioner Alexander MacDonald to see what might be done to save the best shot-dodger of the woods. Out of the conference came the decision of the Conserva- rREFACE lion Uepartnu'iit to inaujiurale a five-year study of the bird, more thorough and complete than any made before. Unlike many surveys, conceived in the necessity of the moment, this one cut its teeth, waxed strong and matured, with the sympathetic backing of sportsmen, into a full-fledged Investigation. Its organization and administration have been the direct responsibility of the senior author of this Report since its inception althouifh. for the most [)art. matters of procedure and policy ha\e been decided in coiisullation with the other authors, who likewise were directly responsible for research into various phases of the broad problem. Thus this report is the result of the comliincd efforts of a group of research workers, at first small in number, later enlarged to include many specialists in particular fields. As time went on. the scope of the Investigation widened. i\ew horizons were brought to \iew and unsuspected fields discovered which had to be cultivated before a harvest of facts could be reaped. The original five-year plan stretched to thirteen years. Though a stopping point has been reached, the end to productive fact finding is not yet in sight. No reasonable amount of time or effort has been spared to make this study comprehen- sive. The work was planned to secure comparable pictures of field conditions throughout the periods of increasing, of maximum, of decreasing and of minimum grouse abundance. Field observations were supplemented by laboratory experiments and examinations wherever nec- essary and possible. In fact, the State's Wildlife Research Centei near Albany was one direct outgrowth of the Grouse Investigation. To allow for regional variations, comparable life history studies covering four widely sep- arated areas in the State were inaugurated in 1931 and carried on. for the most part, to date. The problems to be considered were early divided by the Investigation into ten factors, each possessing the |)ossibilitv of markedly infiuencing grouse abundance. Attention was first focused on obtaining basic life history facts covering food and shelter requirements, repro- ductive capacity, weather relationships and the jiart played by jiredalors. diseases and para- sites, and by man. General habits of the grouse were thoroughly studied, as were the factors limiting the successful artificial propagation of the birds. Only recently there has been added an important study of grouse physiology. All data, no matter how apparently insignificant, were carefully noted, recorded and an- alyzed. Modern statistical methods and machines have been called into play and. through years of constant work, more than a million records have been coded, punched on statistical cards and run througli statistical counting and sorting machines to complete the desired correlati(nis. To avoid ]nissihle })itfalls, each of tiie 186 tables here ])resented has been checked carefully bv the latest biological statistical methods. To carry on the Investigation has been our pleasure as well as our work. How efficiently we have done It. only time can prove. To each generation the efforts of tho.^e preceding often ajjpear ineffectual and misguided. Wildlife research workers and sportsmen in the future, with their more advanced viewpoints and resources, made possible by past successes and failures, may view our efforts in the same light. Indeed, it should be so. AH prowlh is based on going further with being discontented uilii present results. Though, in part, the research has been tedious in detail, as a whole it has bc<'ji iiil<'i('sting and fruitful and often surprising in results, ('ertainly it has given us a new conception of some of the poweifnl forii's wliiili are secretly working under the surface of .Nature. It has PREtACE also brought us to realize how impotent we have been and still are to recognize and con- trol all those forces affecting the abundance of any wildlife species with the knowledge and means at hand. From its inception in 1930 lo the date of publication of this report, one idea has been uppermost — to find ways and means of assuring the future of the ruffed grouse. Far from proving an end in itself, the linestigation has. in reality, opened a gate upon vast and fer- tile fields for future research. It is fortunate that there is always a challenge in the |)Iowinles have become an integral |)art of the wild- life management practices which are helping the Fmpire State to maintain good hunting of several species for some half a million outdoor-minded men. In presenting this xoliime the authors have attempted, in the first chapter, to correlate and preserxe the saga of the ruffed grouse. Leavened with droll notes, for the most part left by the \encral)le describers of another da\ . it will be found to furnish many a jileasant sourre for ((iriicrturc. In the rest of the Miliinic. tlic suimiiai i/.ing nf the findings, and |iarti(U- larly the interpretation of them in semi-lechnical terms in the form of management sugges- tions, must be regarded as tentative rather th;in final. Ortaitd\ thex are open to modification by whatever light ma) be shed u|i(in them li\ future research. If the biologists find, herein, a few new facts clearly presented; if the nature lovers are reminded of pleasant da)s afield; if the sportsmen discover some ])ractical answers to many puzzling questions and the wildlife managers a few helpful suggestions — our own game bag will not hang slack nor will those who ha\e demonstrated an abiding faith in the solution of the grouse problem go unrewarded. Mav 27. 1942 Gardiner Bu.mp. Senior Aullioi 1'^ 9 i >». • tr. • * ; I i •>'; /' y ^r-r-.rt \f .i.f^■^;■ -^^*^ ACKNOWLEDGMENTS "Acknowledgment" is a poor word to fully convey the authors' {eelinp of debt to the many individuals and organizations who have given freely of their time and knowledge, as well as furnishing material aid. in helping to make this book possible. To each and every one we extend our sincere thanks for the understanding and generous coopcratii>ii which has been so vital to the progress and completion of this Heport. May this public recognition, therefore, represent the gratitude we owe them. We regret that each (atmot be cited indi- vidually. Yet we trust that they will fiiul a real measure of reward in the knowledge of hav- ing shared in the undertaking. Preeminent among those to whom we are indebted are those who initiated the Investigation and have been its cooperative backbone ever since. We refer to the sportsmen and to our asso- ciates in the Conservation Department of New York State. \^'ithout George A. Law\er and his committee of s|)ortsmcii. who met at the Department oflices one blistering August day in 1929 to find wa)s and means of helping the "partridge" stage a comeback, the Investigation might never have been initiated. Without the help of a legion of understanding friends both within and without the various sporlsm(>n's organizations, the study coulil ne\er have been carried to maturit). The Investigation was a Conservation I)c|iaitinciil picijccl. Hut siiici' individuals constitute ihe De|)artinciil. the degree of success attained by the project is indeed a measure of the ex- ceptional cooperation which each anil cvcr> one of our associates ga\e the authors tiirough- out the entire course of their work. To the oflicers of the Department, through the years, headed b\ Commissioners Alexander MacDonald. Henr\ Morgenthau. Lithgow Osborne. John A. White, and Ferry B. Duryea. we are especial!) grateful for that inspiration and leader- ship so essential to success. Th<- encouragement and sympathetic interest in the work given us by the Director. William C. Adams, of the Division of Fish and Came, has been unfailing. Throughout the Department cooperation has been complete and whole-hearted. Particular credit is due l)eput\ Commissioners Vial assistance have patiently assisted in gathering and compiling the basic data without which the Report could not have been writ- ten. Theirs was not a spectacular job. Yet the whole web of continuity depended upon its ACKNOWLEDGMENT:^ being vvt-ll (Umw. Id llicni the aiitliors owe a deep personal debt of gratitude. We have !iot met their equal ii] niatn a da). Particular apprei iatiDii is due to Mary K. Thornton who carried on most of the immense task of examiiiiiijr the literature as a basis for compilinfr the bibliography, but who unfor- tunately met wilh a fatal accident before the work was lompleted. In this coimection we wish to thank the staff of the New York State Librar\. as well as those of the libraries of New York City. Cornell I'liiversitN and the American Museum of Natural Histor\ for their co- o|)eratioii in locating references. Few wildlife species have been subject to more stud) than has the ruffed grouse. Vie feel deeply obligated to those who have gone before us for their exploration and delineation of the problem. Outstanding, perhaps, was the American Game Protective Association, whir h has fostered grouse research since its organization in 1912. Beyond these, a number of individuals have contributed generous!) from their experience: have been instrumental in providing facilities for special projects; or have given substantial help in connection with the preparation of the manuscript. Among these may he mentioned J. B. Rurnham. F. C. W'alcott. C. E. Ladd. A. 0. Gross. \^'. L. McAtee. Aldo Leopold. J. C. Phillips. K. T. Frederick. (J. M. Allen. E. L. Brunetl. E. K. Burckmyer. Clarence (;ottam ami .Ario Pardee. Likewise, helpful advice and data have been funushed by the Conservation Departments of Massachusetts. Michigan. Minnesota, Pennsylvania and Wisconsin, by Cornell University, the Massachusetts Fish and Game Protective Association. L'. S. Fish and Wildlife Service and the U. S. Bureau of Animal Industr). Material aid, largel) in the form of labor and materials, has been ct>nlribule(l by the Civil- ian Conservation Corps. Works Progress Administration and State Temporary Emergenc) Relief Administration. For special aid witli Icdmical problems, the authors are also indebted to the following: Fur idciitiliialiori and inter|)relation of food habits material: H. ('•. Harbci. Paul Bartsch, \\ . .1. (!haml)crlain. Adrian Fox, A. B. Gurney. \\ . J. Hamilton Jr.. Carl Ibinrii li. F. J. Hermann. 11. I). House. J. E. Morrison and W. C. Muenscher. as well as tin- cnlirc stall .pf the FikmI Habits SeitioM of the Fish and Wildlife Service. For identification (if parasites: J. C. BiMjuarl, F.. B. ("ram. M. (". Jones. Robert Matheson. R. R. Parker an.l F. E. Wcln. Vav data lorKcrning tlic raiig<' i>f tlii' sjiccics: Rudolph Bemiitt. B. W . (!artw right. C. II. D. Clarke. R. T. King, H. F. Lewis, j. A. Munro. D. 1. Rasmussen. William Rowan. J. I). Soper and L. L. .Snvder, as well as the many reprefentatives of state, provincial and ter- ritorial garni' administrations tlirouglioul the region involved. Fur ln-l|i in -lali-licalK aiiaUziiig llic data gallicred: C. McC. \lolllc\. linalh. ihc autlmrs wish lu |ia\ Iribulc to one who. mcirc ihan am ullicr. has served as ciiunsellor throughout the Investigation — Ur. A. A. Allen of Cornell L'niversity. Deeply in- terested in grouse himself, he has been associated with every important study of the species in the Northeast during the past quarter-centur) . His pioneer attempts to rear grouse in captivity have earned him the a|)pellation "dean of grouse breeders." From such a back- irround. he has constantiv "ivcn ;idvicc and cnconragement to ns. his former students. PART I Thk Ruffed Grouse — Its Background, Basic Biology and Economic Impoutance CHAPTER I THE RUFFED GROUSE IN THE MARCH OF TIME By Gardiner Bump somf-: hkcords of eafily observers first description names by which it is known evidence of high esteem since early colonial times increase in numbers near settlers' clearings reduction in numbers near centers of population the f.ra of exploitation —market hi nt1n(; grouse bounties recognition of periods of scarcity a chronology of suggested causes attf:mpted remedies rise in importance as a sporting bird early efforts at artificial propagation the period of correlated research initiation of tiik i'hksfnt investigation ^ cw Whatever contributes to make our life more attractive usually finds a jjlace in history. So it is that, furnishing both food and sport, the ruffed grouse has reserxed for itself a fascinating niche in man's annals of the past. The larger part of the story is, of course, tucked away in memories, but an astonishing amount has been more permanently preserved in written form. The curtain rises amid the rugged walls of the Frankstown Cave in central Pennsylvania. A group of naturalists led by Alexander Wetmore are huddled around their light expertly sepa- 2 THE RLFFEI) GROISE l.\ THE MARCH OF TIME raling. finrii ihc Ia\iTc(l debris, ihc Ixincs of animals Ion;: cxtiiul. One, small and characlei- istically light, is identified as a part of the cranium of a ruffed prouse. Another is a complete tibio-tarsus. a bone from the iej; of the same species. W ilh these are uncovered parts from tlic skeleton of a turke\. The scene shifts to the Cumberland ('ave, broken into while makiiin readily available and to stimulate interest in and thoughtful consideration of it. the more informative records have been coUeited for this chapter. Many of the quaint colloquialisms and the naive expres- sions found in early writings have been here reproduced by way of piquing the curiosity atid seasoning the Report. SOME RECORDS OF EARLY OBSERVERS Habit and size decreed for the ruffed grouse a place of importance among our native birds. History has. accordingly, been generous with her records. One surmises, from the broken grouse bones that are occasionally excavated from their camping grounds, that Indians found the birds to their liking. But it is with the white man that the story really begins. In L531, to his roval household in Kent. King Henr\ Vlll issued a decree controlling the shooting of "grows"'. The name, perhaps anglici/cd from the old Erench liriesch. meaning spotted, had been applied, not only to the introduced blackcock for which the decree was issued, but also to its much more common cousin, the red grouse. Arriving in the New World. French and English colonists. carr\ing with them impressions of Old World grouse, promptly named the spotted bird whiih haunted those woodlands, grouse or partridge. Farther south, it became the moimtain pheasant, a name by which it is still known among the back country folk of today. FIRST DESCRIPTION 3 As early as 1632, Thomas Morloir"", early advcnlurer and Indian trader, records seeing 40 "rouse in a single tree, an indication of their abundance at that time. Later Nicholas Denys", French representative in Nova Scotia, in 1672. reports their tails were used as fans in France. He tried twice to transport the birds there, "but when approaching France, they die; which has made me believe that our air must be contrary to their good." Despite the tendency for early observers to confuse the spruce grouse with our ruffed grouse, it seems fairly certain that Baron de Lahontan"' was referring to the latter when in 1703, in his "Some New Voyages to America", he comments on the abundance and stupidity of what he called the woods hen, "for they sat upon the trees in flocks and were killed one after another without offering to stir", writes the Baron, who added that this phenomenon was the most comical thing he had seen in America. Less sure are we that Abbott' was not confusing this si)e(ies with the eastern heath hen, when he indicated that, in the closing years of the eighteenth century "these birds used to con- gregate by the thousands" in the swamps of central New Jersey. Here it was common sport to surround them and either shoot or net them in huge numbers. That the grouse unwittingly pla\cd still other parts in the lives of the early settlers is gathered from many a record. One unusual instance will suffice. Rumor had early linked deposits of desirable metals— lead and iron— with New York's Adirondack Mountains'". His vessel becalmed one warm day near Red Rocks on Lake Champlain. ski|)per A. M. Martin crooked his arm around his gun and sallied forth in quest of partridges and a change of diet. Following a bird that had flushed, he tried to pull himself up over a small ledge by grabbing a small bush. When this pulled loose, it uncovered a dark, interesting-looking stone. Stick- ing it into his pocket, he hurried on. Later he sent it to Ticonderoga. there to be recognized as high-grade iron ore. Thus were the Penfield iron deposits discovered. FIRST DESCRIPTION Explorers and the colonists that followed ihcni were, for the most part, men "f action and of the land. Few among them were interested in the natural sciences. It is not surprising, therefore, to find that the grouse, in common with many another bird, did not receive taxono- mic recognition until a century and a (juarlcr had passed. John Bartram. an enthusiastic traveler, sent a letter to England, in 1 7.S0. recounting some habits of the bird. This was accompanied by a specimen which was described In Edwards, a British naturalist" . This eventually became the type specimen because on the basis of this account. Liimaeus included the species in his Systema Naturae™ which has since been designated as the foundation of modern nomenclature. NAMES BY WHICH IT IS KNOWN What's in a name? That which we (all a rose Bv any other name would smell as sweet. Shakespeare So it is with our native ruffed grouse. This bird carries such a myriad of names so diverse in meaning that a casual glance at a list of them might create the impression it represented several species, rather than one. The unwary "fool hen"* of wilderness areas is the alert "king of game birds" where heavily hunted. The "pine hen" of the west, the "birch partridge" of the Northeast and the "spruce-woods rufliled grouse" of our northern climate are one and * ^..^v iiinr.- roiiiIiMniU upplicd to the spruce protise. 1. Tilt: in I FED (,ll<)LSI: l\ HIE MARCH OE TIME ihf same witli tlic "pats'" i>f Midiijian and tlic "mountain cock" or "plicasanl of the Soutli. The diversity of these pieturesqiif cliaraMs is so great and their contrast, in many cases, so striking, that it may be of interest to look into ihe maimer in whii h liiey probably originated. Indians |)icturesquelv referred to it as "larpenter bird" because it "pounded on a log""'". Linnaeus, in first describing it, assigned it to the genus Telrao with the Kuropean grouse and specifically termed it umhellus because of the umbrella-like ruff which, when erect, framed the head. This also accounted for the most commoidy used name, "rutTed grouse". Later, however, the generic name Boiiasa ( Lat. bonasus = bison ) was used, likening the drumming of the grouse to the bellowing of a bull. Thus, scientificall) . the ruffed grouse became Bonasa uinbeUus. The word "grouse" seems to have been derived frcjm the French terms greorlie, ^reiclie and priais, meaning spotted bird, and in the form "grous" was used in England before being applied to New World species. Among the Indians and Eskimos, its names were as many as the dialects of the varying tribes which frequented the range of the bird. No record remaitis of most of these. Then came the white man: Spaniard. Frenchman. Englishman. Scandinavian. German and the rest. As .settlement proceeded, these various races tended to colonize in different sections. It was quite natural for them to appl) to the new birds the\ encountered, the same names used in their homeland for species of similar appearance. Memories of the blackcock, the Scotch grcjuse and of the ancestors of our adopted ringneck pheasant, gave rise among the English to such terms as "moor-fowl", "ruffled heathcock"'. and "mountain pheasant." Similarly in French Canada, the ruffed grouse became "La Grosse. Gelinote de Canada"' and "Coq de Bruyere a Fraise", while the Pennsylvania Dutch likened it to the German "fesond." In addition to names derived from the racial background of the early settlers, many others have had their origin in distinctive characteristics or habits of the bird. Those mentioned above do not need further explanation but a few more will round out the discussion. The nature of the meat suggested the designation "white-flesher"". while its unique drumming earned the name "drunnner" in many localities. Thoughout the Northeast, where birches were a favorite food, it is commonly known as the "birch partridge". At the other end of its range, how- ever, Ozark mountaineers call it "woods pheasant" as di.stinguishcd from the liobwliile quail which is still known as "partridge" in the South. Remembering that the ruffed grouse has adajjted itself to half a continent, it is not surpris- ing that so man\ dissimilar and localized Tiames appear in the literature. Dniilitlcss there are niaiiN more lh;ui listed here wliicli arc localK conunon but i>f which no record has been found. NAMES BY WHICH IT IS KNOWN Names Used in the Literature in Referring to the Ruffed Grouse Bv non-English speaking races Indian names bi-ne (grouse in general) carpenter bird ka-xit'1-se kh-tuk (Eskimo) kwi-ut -- pah-uta pupuskee puskee wen gi-da-bi-iie (Chippewa! French gelinotte greoche griais griesche English grous moor fowl more-hen ruffled heathcotk Canadian liirch partridge I Br. ProNiiues) Canada ruffed jirouse Canadian ruffed grouse French hens I Labrador and Quebec) patridgp (Maritime Provinces) shoulder-knot grouse white-flesher while-nieated grouse willow grouse (Br. Col.) Northeastern L niled Stales American partridge birch ])artridgp (Maine) comnion partridge drunnning grouse (New York) eastern ruffed grouse (connnon) partridge (conunon) palridge (common) ruffed grouse (common) wood-pile quawker (Long IsLind ) Fren ch -Canadian attagen Pennsylvaniae coq de bruyere a fraise grosse gelinotte du Canada la gelinot hupee de Pensilvania la grosse gelinotte de Canada la perdix ordinaire perdrix franc German fesond haselhuhn Scandinavian hjerpe By English-speaking races Southern United Stales moore fowl (So. Carolina) inounlain cock (Carolinas) inoiinlain grous (Carolinas) riMiuiitaiii partridge (Virginia) rnouiitaiii pheasant (Virginias and Carolinas) pheasant (common ) ruffed grouse (common) Western United Stales fool hen (Rocky Mts.) Oregon grouse pats (Michigan and New England) pine hen ( Vi asatch Mis.) .^abincs grouse western ruffed grouse wood pheasant woods pheasant (Ozarks — Missouri) Tilt: Hi I FEU GRUL >'£ l\ THE MARCH UE TIME Color lnown riifled trrouse copper ruflcd grouse gray mountain grousp gray ruffed grouse gray ruftled grouse gray-tail red-ruffed grouse red-tail silver-tail Based on sirurture ruffer rulTet! grouse li|)pet grouse Based on sul)-species according to: Distribution Arctic ruffed grouse hush pheasant Canadian ruffed grouse eastern ruffed grouse northern ruffed grouse Oregon grouse western ruffed grouse Based on habits drummer drumming grouse druniniing partridge (Irumining pheasant fool hen ruffled grouse EVIDENCE OF HIGH ESTEEM SINCE EARLY COLONIAL TIMES That the "fool hen" characteristic of the bird in the early days did not affect the high esteem in which it was held as a piece de resistance for the table, is clear in all r)f the early writings. A letter from John Bartrani. sent to England about 1752 and quoted iiy Edwards, states, "their flesh is white and good". Audubon opines that the grouse "far surpass, as an article of food, every other land bird except the wild turke\". while Billings'", writing at the same time, reaches a similar conclusion. Larger than the quail and more abundant than ihc tiirke\ tiirouglioul tlic northeast, half a century ago. one might have found grouse competing with ducks in po|)ularit\ in the mar- kets of the larger cities and definitely more common in rural kitchens. Tiiat todav it can rarely lie shot on the ground over mucii of its range, lias, of course, made grouse a less common but more prized delicac\. to which the auliiiiiM-bniw ii of wooded hillsides, leavened with falling lea\fs and tired musc-les. often adds a (liquant llaxor. With more hunters than ever afield, turning their attention in increasing minibers from pheasant to grouse, the enthu- siasm for the sport vies with ils thrills in |)utting the ruffed grouse in a class by itself. For. to be a good grouse hunter, with dog and gun or a camera, is to |)lace one's self in a rather select circle of outdof>r devotees. Thus time has ser\c(l to >hifl llic rrnplKisi> licini grouse as an article of food lo a place of high favor as one of the most >porling birds thai enriches our co\erts. INCREASE IN NUMBERS NEAR SETTLERS' CLEARINGS Lcgendar\ have become the laics of llic iniiltiludc of grion of llicii < rude clearings and cuttings was. in a large measure, the cause of this conc<"nlraliori. ."^wainson and Richardson'^ slate that grouse "frequents the horse-palhs and clearings about the forts', while Alexander Wilson"" records that he could always snpplv himself with plenty of birds "without leaving the path". REDUCTION IN NUMBERS NEAR CENTERS OF POPULATION 7 Today the effect of woods edges on the abundance of many a wild-life species is well rec- ognized. Perhaps more woods-wise but certainly less analytical, the early settlers utilized this source of fresh meat close at hand. They seldom questioned (if we may believe the records) but that the woodland beyond the ring i>i their axes likewise held countless thousands of birds. For then, unlike now, the hunter apparently rarely inquired as to the cause of such concen- trations. They preferred rather to harvest the crop and. when the number of grouse in their coverts periodically shrank to a fraction of their former abundance, to hope for better days. Thus the axe. where not followed too closely In fire and the plow, served to break up the forest cover and. as with deer, to increase its carrying capacity for game. Partridge thrived accordingly. REDUCTION IN NUMBERS NE.AR CENTERS OF POPULATION The diffprence between optinuim and poor grouse habitat is often a matter of the degree to which forest cutting and clearing are carried. Men were eager to turn the forest duff into fertile fields. It was. of course, inevitable that, aniund the larger centers of population, where this activity was greatest, food and shelter for grouse became progressively less and hunters more numerous. This led to a gradual reduction of the species in such areas. Bartram '. about 1752, noted the disappearance of the grouse in the lower settlements of Pennsylvania. Sixty years later. Alexander Wilson"" saw the same trend and wrote: "formerly they were numerous in the immediate vicinity of Philadelphia but. as the woods were cleared and the jjopulation increased, they retreated to the interior". Some of this range, in the East, has Ix-cn taken over by the pheasant and the bobwhilc quail. All too nnuh of it. particularK uhere it is heavih |)astured has been left barren of any game bird, save possibly the woodcock. b\ the retreating feet of the jiarlridge. Influences generalh little recognized, in the form of the machine and of scientific agricul- ture ha\e recently made farming unprofitable in so much of the hill land and back country that once was grouse habitat, thai the trend has been reversed again. Woodlots are going lirush). and woods" edges are growing up to a variety of plant species furnishing both food and shelter for the grouse. In New York alone, upwards of r).0()().O(M) acres of once-farmed fields are now in process of abaridonment. Even the lands of medium quality are apt to be worked less intensively. The woods are accordingK. growing back toward the centers of population, bringing the grouse in tlieir train. .^$>X*A-.- THE RL Ft ED GROiSE l.\ THE MARCH OF Tl ME THE ERA OF EXPLOITATION— MARKET HINTTXG The use of the grouse as a source of food for the home tal)le li\ the early settlers was followed naturally by the exploitation of the species for the market as commercial facilities developed. The price which these birds brought in Boston. New York, and Philadelphia, where most of them were shipi)ed. ranged from 12^ ^ cents to .'?2.()(t a])iecp. depending upon the abundance and proximity of the supply to the place of sale. The all-time high is a recent record in 1918 of .S7.00 a pair in Washington. A few of the more interesting records are as follows : Place New York Philadelphia Boston Boston Cincinnati Eastern Cities Pittsburgh Kingston. N. \. Maine New York Maine New York Chicago New York City New York City Price List St. Stephens. N'. R. I Canada I New York Michigan Washington. I). C. Date 1763 Around 1812 1820 1820 Late 1820V About 1825 1877 1 !!86 1890 About 1894 1899 1902 1902 1908 1908 1910 1911 Market Price .24 each .7.i- 1.00 each .12M > each .50 each .121: ; each .7.5- 1.00 each -12M > each .40 brace .08 each 1..50- 2.00 each 1.00- 2.(H) pair 2.00 pair 10.00 doz. 1.25 each 1 .00 each .60 each 1.7.5- 2.00 each 1.0(1 each 7.00 pair 1918 Market hunting was a profitable occupation througlioul nuiili of the grouse range, largely because of the ease with which the bird, when occurring in sudicicnl nmnbers. could be secured. Methods of bagging them for market were as varied as the ingcnuil) of the hunters. Snares, lures, traps and guns all pni\c(l rlTcctixr. Tun of llic iiinii' uni(]ue means THE ERA OF EXPLOITATION— MARKET HUNTING 9 are described by Nuttall""". One is the imitation of drumming to lure '"jealous males" within shooting distance by striking upon an inflated bladder. Another is the smoking to death of the birds "in the same manner as the wild pigeons in the western country." SNARES CONFISCATED BY THE NEW YORK STATE CONSERVATION DEPARTMENT A very amusing and interesting story cm tra|.i)iMg. illustrative of the general practices of the times, is given by Whitehead when writing in Mayer's'^ "Sport with Rod and Gun." as follows: "This bird is the friend of the country boy. It has many a time made him jump as it l)urst out of the wayside bushes, and bird and boy perpetually match their wits against each other —the one in trapping and the other in avoiding being trapped. Master Barefoot finds a drum- ming log, and at once whips out his jack-knife and. bending down a neighboring hickory sapling, sets a twitch-up. with a slip-noose at the end. made of a string pulled out of one of his capacious pockets. The twitch-up being well watched, is sure to catch the bird or drive it away. As Barefoot grows older, he learns to set running snares of horsehair or silk in the paths in the woods, and he will walk miles to attend them when he is too sick to go half a mile to school. At length, he grows to be a young man, 'some farmer, some poacher', making a precarious living by selling game he has trapped or shot in season and out. and killing more birds than all the minks, owls, and foxes in the countryside." A method, formerly common and still practiced illegally in remote parts of New York State, is to construct one or more low brush fences running through a cover frequented by grouse. At intervals, openings are left in these fences, the ground in the immediate vicinity being cleared of leaves and other debris to attract the attention of the birds. At each break, a wire snare is so set as to be sprung by a grouse walking through it. Once released, a springy stick promptly jerks the unfortunate bird by the neck, several feet off the ground. Grouse are prone to do more walking than is generally realized. A brush fence, therefore, is an effective means of guiding them to the openings in which the snares are set. 10 THE RUFFED GROUSE IN THE MARCH OF TIME One can come closer to wiping out these birds, in a patch of woodland, with snares than by any other means. Edward Howe Forbush"" for many years state ornithologist for Mas- sachusetts, reported ''two men in the vicinity of Westfield, Massachusetts, took one hundred and twenty of these birds from snares in one day". Five men of Foxborough snared grouse prior ti) 1888 and averaged about a hundred birds a week. The most successful snarers probably never published their records, for obvious reasons, though one finds, in an 1878 issue of "Forest and Stream", a notation that 1,189 grouse were snared in 54 days, an average of better than 22 a day. We have seen somewhat similar arrangements designed to catch grouse by the feet at their dusting places. Fox traps set at the edge of spring-holes in winter still take a surprising toll of feathered game, a method doubtless also employed by some marketeers during the off-season for other trapping operations. But shooting remained the most popular method wherever large numbers had to be secured at definite intervals. Unlike today, around every center of populati(m there was always plenty of back country into which but few of those interested in grouse, save the market hunter, pen- etrated. There grouse were undisturbed and so were comparatively unafraid of man. For- bush'" quotes E. F. Staples of Taunton, Mass, who writes, "This gentleman said that, in the early 1880's. about a thousand birds were killed in a season on 20.000 acres that he ranged." It was not unusual, in a good year, for one hunter to kill from 30 to 50 birds in a day. Careful inquiry, however, indicates that the average was substantially lower than this. Mershon"', when a boy. found ten to twelve birds a day to be a fair average under Michigan conditions. Also, Maynard in 1870"°. states that market hunters from Worcester, Mass., killed ten to fifteen birds a day. While the year's best hunts, then as now, were long remembered, personal contact with several of the fast vanishing market hunters in the Northeast indicates the aver- age daily bag in this region to have been much higher. Most birds were shot on the ground, an item seldom ap|)reciated when one compares tlie take of yesteryear with today's much smaller legal limits. Roads, trails and woods edges were favorite places, for there the cover was open and the birds more apt to be concentrated, partic- ularly in the summer and fall. When spring shooting was permitted, drumming logs often provided an unexpected rendezvous with death. So highly commercialized were these activities that individual hotels, in many a large city, employed their own hunters. One trapper in Chenango Countv°' reported selling over 3,500 grouse to one New York hotel. Some 2.500 birds were shipped to the New York City markets from the Poughkeepsie railroad station alone in 1875". Mr. George Howes killed and mar- keted 398 birds in one shooting season while Haight'" relates the fact of two men taking 998 grouse from Septeinl)er 15 to January 31. »ne season in the late 1800s. Down from the timbered hills to the markets came the grouse bv the hundreds of thousands — in barrels, in boxes, in bags, by horse or traiTi — until the turn of the present century. Almost from the beginning, a few men had decried the [iractice. \uttall"'. though (irima- rily a botanist, was also a keen observer of animal life. As earlv as 1r('rmber 26. 1911. 12 THE RIFFED GROUSE I\ THE MARCH OF TIME so unusually abundant previous to that period as to sell in the markets of Boston for as low as 12*2 cents each". Of more than coincidental significance, therefore, is an editorial carried by the Rochester (N.Y.) Dailv Advertiser of April 11. 1832, beginning. "\^'e have regretted the destruction of this bird by the hard winter of 1830-31, because his note was necessary to the chorus of the field and the forest — the only chorus that we ever admired", and urging its fur- ther protection. The significance of this evidence of scarcity during the same year from sources 600 miles apart, was to remain unrealized, however, for most of the next hundred years. But not so with the records of scarcity. A letter from a market hunter of the old school, Mr. W. E. Hookway of Syracuse, N. Y., to the authors, states that a companion of his father's, engaged by the old Saratoga Hotel, killed only one grouse in five months during 1867. As though to corroborate this observation for other counties of the State, a few years after the Civil War, Judge John W. Spoare, a locally famous grouse shot of Columbia County (N.Y.), is reputed to have sold his only hunting dog because "the partridge are all gone". That coming events sometimes cast their shadows before, is indicated by Norris^, who records grouse as common in Massachusetts but scarce in Vermont in 1866. "The Fluctuating Grouse Supplv"' is the title of an article published in Field and Stream in 1883. According to the author, in southwestern Ontario, grouse were reportedly scarce, in shar]) contrast to their abundance two years previous. Apparently reliable though fragmentary reports indicate a period of scarcity in 1896 or 1897. In substantiation of this. Dillin"" records that, in two weeks, he flushed two birds and his companion but three, in southern PennsvKaiiia in 1897. whereas they had averaged 14 per day in 1889, on the same area. With the turn of the century, interest grew apace. The first comprehensive attempt to outline the problem followed the exceedingly severe winter of 1903-04. when Edward Howe For- bush'" sent a questionnaire to some 400 interested naturalists, game wardens and sportsmen throughout Massachusetts, inquiring as to its effects on bird life. Grouse, as well as other wild birds, were reported exceedingly scarce. On the u inter rested most of the blame. The scarcity of 1906-07 was unique in that it was so generalK recognized by observers. Writers recorded that the species fell to rock bottom and it was frankly ])rcdicted that the end (if the grouse was at hand. The New York Forest. Fish and Game Commission became so alarmed that an extensive survey by questionnaire was conducted to "bring out" the situa- tion (Woodruff'"*). Attention by now was aroused to the rather regular recurrence of these periods. When the next "die-off" occurred. Stoddart'"'. then Rod and Gun editor on the New York Times, was ac- cordingly engaged by the New York Commission to circulate an even more extensive ques- tionnaire, particularly among hunters and game protectors, to determine supposed causes of the lean years of 191.5-16 and 17. Now, too, the American Game Protective Association, formed in 1912. was becoming intensely interested, not only in attempting to propagate grouse artificially, but also in the causes of their scarcity. An off-cycle decrease in 1924, throughout much of the grouse range, was sufficiently intense to cause the formation of the Grouse Investigation Committee at the American Game Conference that year. Less than twelve months later, throughout the North and Middle W est. competent observers, Leopold"", reported ruffed grouse to be at the numerically low point of another cycle. Throughout the East the tide of scarcity reversed itself, surging upward for two short years, then shrank to the well remembered low of 1927-28. A CHRONOLOGY OF SUGGESTED CAUSES 13 By now everyone was cycle-conscious. The present Investigation was born of this scarcity in 1930. Following its inception, the birds were reported as having practically disappeared in the Northwest and far North in 193.5. As though to make the work more difficult and challeng- ing, an anticipated scarcity, in 1937, throughout the East, did not materiahze though the numbers of birds were not up to par in 1936 in New York State. A CHRONOLOGY OF SUGGESTED CAUSES From the earliest records, the reasons for each period of starcil) has been conjectured by its historians. Some, usually interpreting limited experience, have oifered a single, simple solution. Others, of a more imaginative turn of mind, have submitted numbers of potential explanations. The more important of these are listed here in table 1. Bartram", perhaps comes closer to the truth than many when he attributes the disappear- ance of grouse in the "lower settlements of Pennsylvania", to the encroachment of civilization. From that early date most of the recorded causes, though legion, may easily be catalogued under one or more of the four "principal changes" su|)posed to beset grouse. In addition to hunting, already mentioned, there were predators, bad weather and disease. Thus hunting did not remain the only recorded cause for long for a scant six years later, a Mr. Brooke of Maryland, in a letter to Edwards'", complained that "the great number and variety of hawks in Maryland feeding on them (grouse) prevents them from increasing fast". This emphasis on the effect of possible predators has held the stage as a favorite cause of decimation ever since. Nuttall took an easy "out" in explaining their disappearance in New Hampshire, in 1831, by deciding that they must have migrated. It is probable that, being somewhat of a cataloguer, he was influenced by Audubon's earlier reference to migration. Already referred to is the newsy comment in the Rochester Daily Advertiser ascribing the shortage in the partridge crop to the severe winter of 1830-31. Local explanations were, of course, rife after each disappearance. Dillin"" adds a new note, attributing their disappearance in Pennsylvania, in 1897. to forest fires. The circum- stances surrounding the issuing of Forbush's qiicslioiinaire into the effect of the severe winter of 1903-04, may have strongly influenced the verdict against the extreme weather, the effect of which was under investigation. With the phenomenon now better recognized, logical explanations became more difficult. In- dividual opinions were less well received and questionnaires, therefore, came into their own as a method of bringing the collective judgment of men of wide experience in the out-of- doors to bear on the problem. Thus, when 1904's lean year, in certain regions, followed by a partial recovery in 190.5. was in turn climaxed by a die-off of even greater proportions in 1906-07, New York's Forest. Fish and Game Commission employed E. Seymour Woodruff'"* to 14 THE RUFFED GROUSE IN THE MARCH OF TIME C/3 ■< O c« O o o w 1/3 o w z a o e o <>: a ca 3q r W O eft >- J O o o z o c: u n 9nuQug ea 6j»iuni{ ui sseaaau] a ■o eaig leojoj ::: :x :::::::: X :: :x :x :::::;:::: : a ijode joj 8uiiooqBJ3A0 : : : : :xl :x : : : : : : :x :j-! :x : : : : :o 3fuiiunq loiiJB^ ixx : : :x :XO :xx :o : :xxx : : : :x : : :X i I S9)I6BJDd |BUJ9)X^ ;><^ iX ix :::;:;:;: :X aeeaeip jo oiuiapid^ ■.X :o ^oiui jo Xipjeog 9aj«q pue e)tqqBj jo X)i3jB3g sifMBqsos) ;o :x ;x :x! suMsq uiBjjBQ X ::::::::::: : : : : :x '.X siMo UIB1J03 ::;::::::;::: : : : :x :>< :o : s3uidn3jo^ ;xx SIB3 siuniS SOXOJ ■.X :oXX :oox : IBJBUaS Ul SJO)Bp3J(J :xx :x :x ;x •.X g la o 2 u 2 :::;;:::::::: ::: :x :X ::::::: : ! :i< uiqBjoABjun uoBB3fi Suip^ajg ::::::::::::: o :x :XXo ixxxo :x : : :X ja[|)e3M pBg :X :X a)Bjdiiu o^ inui^suj I '^X '. I * ' ! ! !X poOJ J»)UIM Ul a6B3J3aQ pooj ^uapmnsni :x :x 183JOJ jjo ^o au^nQ :X :x :X :X JSAOO d|qi^!nsu|^ Q (S c c cc=eec >-H >t- > > >>>>.t ') c a B c = =ov o 3 o;=o:s 2 Z iSZZiCZZ 0 0 0 0 Z222 o c > 22 £ a. wwp3^iftwoor-r-r-r-oecccoooo>ovo COCOCOCCOCOOOOOCO-N 111 11= *3 oSa ■2-J i< si" j< >j= = =-J I? 5i-= -^ ?^.= =■ A CHRONOLOGY OF SUGGESTED CAUSES 15 H O u o a; O g • O t/D "^ "I ^5 z a o o -j2 ■<^ ot O Q :« W o z o « c ca a •c £ a 1 Suueug : ;x : : : : BJ9innq ui dseajao] X : : : : :o : : gajlj 183JOJ : :: :x : : : : X : : : : : : : ;x ; Ijode joj ani^ooqejaAO ; :x :o : : : X : : : : : :o : : : aojiunq ^a^JBiv X :oo : ; ; : : :o : : :x : : : : s 1 .ill Q g es^isBJBd fouja^x^ ;x : : : ; ;o : :o : : : ; oseasip JO otaiitpul^ : : : :x : : : ox : : : XXO : : :x o :o E 2 ffl aoiui JO XipJBOg X : : : : ; ; :X sajeq puo n'qq^j JO Xiiojbos : : : : : :o : xo : :x :ox :o : BUMBqeoQ : : : : : :x : X : :o oxo : :x : si(MBq unrija3 X : : : : : : :X :xx : : : :x : eiMO UIB)J33 : : : : : :x : X : : : : :x :xx ; : ; Bauidnajoj «10D : :x : : : : : fiHuniis X : :xx : : : Baxoj X : :xx ; : ; IBjauaS ui sjo^npajj : : : : ixx ; XXX : : : :x :xx :ox 1 i jaqinoM ppo X :X aiqBJOABjnn uosBas Suipaajd : : : :x :x : X : jaqiB3M pBQ :::::::: :XX : X : : : OlBjailU O) 13U|-)SU| : : :x : : : 1.1 fees pooj ja)uiM III asBajdaQ :x : pooj jiiaiaiunsui :x : isajoj jo jjo auiiin^ : :x : : : : : :x : : : : ;x : : : jaAoa a|qB|in8UQ :x : : :x : : : : >> " Si Not given Not given Not given Not given 1915-1917 Not given 1916 Not given 1917. 1924- 1926-1927 1916-1917 Not given 1918 1918 1918 1918-1920 Not given Not given 1925 1926-1927 1927-1929 Not given Not given 1928 1928 Not given Not given Not given Not given DaU of publi- cation 1914 1914 1915 1915 1923 1916 1918 1917 1924 1929 1918 1918 1918 1918 1921 1922 1919 1923 1925 1928 1930 1927 1928 1931 1928 1930 1932 1932 1933 1 3 < Rough. J Hodge. Dr. C. F.. Ridgway. R Reiner. N. E JackBon. H. H.... Field. Geo. W.... Burnham. J. B. . . Scudder, B. A. . . . Allen. A. A S : ': IP llj Mousley, H Bradshaw. K Editor Corson, G. 11 Uross, A. u Sutton, (J. M Bump. G Bent. A. C Huntington. D. W Luttringer, L. A.. Anonymous Roberts. T. S Laing. H.M Van Ness. Vernon l&^ .= «- = a o ; 11 ■="2 M. IB ?•,__ g|i"o -•- >-2 — 00 m an ■=°ai 0 8^2 _go-oo =•11 § E o >.S 1*1° I -11 = o-2 8:r2 C£ II i 3 S.- .S 3 " * 1 " a Ei Si"" £ s .2 gi- » . bfi§ a S'a« "3-3 3 2^ .2 » » .= — 13 *- « . oSgg-2 0 " S- S .= .. u es 3 3 - _ u b£ V o 5 Ca 3 16 THE RUFFED GROUSE /A' THE MARCH OF TIME condui t a survey, by questi(jiinaire, of the probable causes. The answers ran the gamut of possible influences, with the fdllciwiiig nine theories predominating: 1 . The severe winter of 1906-07 2. Unusual abundance of foxes and goshawks 3. The cold wet spring of 1907 4. The extreme dryness of the following July and August 5. A disease epidemic 6. Internal parasites 7. An infestation of parasitic ticks 8. Pot hunting in the closed season 9. Resumption of migratory instincts With commendable inclusiveness he concluded the best bet to be an unhappy combination of the cold wet spring, the unusual abundance of predators, and an epidemic of some disease or parasite. By now, interest was general and the outdoor magazines of the day were filled with causative speculations. Biologists, too, were sharing in this interest. Thus George Bird Grinneir% in 1910, lists six of Woodruff's nine factors as suggestions but places the most emphasis on the cutting off of the forest, with over-shooting as a close second in importance. He was also the first to publicize the periodicity of the scarcity phenomenon. In the meantime, striking progress was being made in raising and keeping grouse in captiv- ity. Poison had brought an end to one of Dr. Hodge's pioneer attempts at Worcester, Mass. as described later. Encouraged by the newly formed American Game Protective Association, Torrey, Dimmick and Fields, in Massachusetts and Walcott and Mac Vickers, in Connecticut, each watched and described the psychology, the food habits and the general reactions of the bird in captivity. In 1915, John Burnham arranged for George Jeffries"" to pen grouse under natural conditions, feed them natural foods and raise a few young grouse to maturity, in an attempt to discover causes for their fluctuating numbers. All this contributed lo an understanding of the bird but did not soke the problem. Inter- est remained intense, but. with magnificent disdain for the efforts of the investigators, the grouse calmly proceeded to disappear again in 1916-17. The distractions of war notwith- standing. New York's Conservation Commission pn)ni|)tly organized a survey under the direction of A. M. Stoddarl which was more far-flung than any >('t attempted. Game protec- tors were ordered to submit weekly reports covering their field ()lis('r\ ations of grouse abund- ance and habits. The presence of some birds, dead in the field, was noted, but .surmise was still substituted for the more certain results of scientific examiiiatidii to (Icicrminc the cause of death. The methods, basic to good wildlife research, were as yet largely unrecognized. While at- tem|)ting to get at the facts thrcnif;!) the directed iibs<'i vatiori of man) individuals, Stoddart was, accordingly, forced to fall back on the most extensive (picslioiniaire yet devised to furnish the broadest possible backgrountl for his explorations. His report, as finally brought togeth- er, leans strongly toward predators as most likely to be responsible for periodic scarcities. These are listed in the following order: hawks, owls, weasels, foxes and the common cat. To this list were added: disease, the rigors of winter, slaughter by man. and bad nesting seasons ATTEMPTED REMEDIES 17 —none of which were new to the picture. An all-time high was reached when he listed thirty- six additional possibilities. A few of the most choice ones are mentioned here for the reader's delectation : driven out by pheasants, shooting from autos, summer boarders, foreigners with- out licenses, and too long a rabbit season. ■ ^ Another of Stoddart's list of causes, entitled "new stock needed", is the hrst indication of the suggestion, common in the late 1920s, that the bugaboo of inbreeding might be at the root of the trouble. Thus was completed such an excellent job of canvassing miscellaneous opinions that no further inspirations have come to light, at least until the period of modern investigation. ATTEMPTED REMEDIES Conjecture as to the factors possibly responsible for decimating the grouse was and still remains a very popular occupation. To translate even a few of these supposed causes into preventative practices proved to be quite another matter. Of course, the first remedy tried was to control the hunting of grouse. It is surprising to note how early this was applied, for a closed season on ruffed grouse, heath hen, quail and wild turkey, was first promulgated in the Province of New York in 1708""'. This was soon followed, in 1721, in Quebec, by an "ordinance to prevent the destruction of Partridge""". The American Revolution brought home rule— and problems, including game ones — for in 1791. the hunting of partridges from April 1 to October 5 was prohibited in three counties now a part of New York City. Thus the evils of spring and summer shooting were first controlled over 150 years ago. The temptation to shoot partridges for sale in the closed season was so strong that, in 18.37, a law prohibiting their purchase or sale in four counties, was passed. The application of this law was extended to the rest of New York State in 1862. Again, in 1839, a closed season was voted to include the whole State"". Other states, too, resorted to similar measures as the best way. then known, to bring the grouse back, as is evidenced by Massachusetts' action in 1818, and Ohio's in 1857. Other controls followed in time. In 1903. spring shooting was abolished in New York and market hunting was legally outlawed. Many other northeastern states took similar action at 18 THE RUFFED GROUSE IN THE MARCH OF TIME about the same time. Shorter open seasons were firmly established and made efiFective through proper protection, backed up by hunter cooperation. Few further changes have been made since, except to vary the length of the closed season, as in 1928-29 in New York State, when no open season at all was permitted. While the protection thus granted certainly helped, it did not solve the problem of periodic disappearances. Interested sportsmen, therefore, tried other methods. Grouse were imported and released on Staten Island (N. Y.) about 1880. to give the few birds left there the added impetus of numbers. Outram Bangs reported a small shipment of birds from north- ern New Hampshire to have been liberated by him near Wareham, Massachusetts, in the lat- ter 80's*''. Bureau of Biological Survey records indicate a shipment of 56 Canadian birds was TABLE 2. RECORDS AND RESULTS OF GROUSE LIBERATIONS KNOWN TO THE INVESTIGATION Date Number liberated Source Wild or reared stock Where liberated By whom Banded or marked Results 1880 30 ± Small shipment S6 38 or .-i* 400 8 of 30 purchased ? 72 115 of 126 purchased 72 A few A few 100 100 A few Wild Wild Wild Wild Wild Wild Wild Wild Wild Wild Wild Staten Is., N. Y. Wareham. Mass. Green Bay. \\'is. Anticosti Island Westchester Co.. N. Y. Connecticut Island in Puget SouikI Westchester Co.. N. Y. Connecticut Westchester Co.. N. Y. Fall No. New Hampshire Canada Nova Scotia Alberta AlberU Alberta Alberta Alberta Alborta Alberta Outram Hangs \N'illlam Bartihard 1907-I*J08 19 survived the first winter: bred and spread 1921\ 1922/ So. N. Y. Fish & Game Ass'n Sturgis & Taintor Marked 1921 So. N. Y. Fish & Game Ass'n Sturgis & Taintor So. N. Y. Fish & Game Ass'n Pa. Game Commi»4it>n Marked 1921 Marked 192-1 Maryland Weatclu-aler Co.. N. Y. Putnam Co.. N.Y. Cypress Hills. Saskatchewan Martha's Vineyard C'onii. Hill Pharsalia (2oiui. Hill Cnpt. Dist. (Jame Hef Pharsalia Came Hef. Conn. Hill 192S Alberta Alberta Saskatchewan Alberta Alberta Alberta N. Y. S. Uhacii Game Farm Alberta Alberta Alborta Wild Wild Wild Wild Wild Wild Reared Wild Wild Wild So. N. Y. Fish & Game Ass'n Noble Marked 1925 1930(P) Feb. 1931 Feb. 1931 Dec. 1931 Dec. 1931 Nov.-Dec. 1931 Mass. Fish *V (iame Div N. Y. S. N. Y. S. N. Y.S. N. Y. S. N. Y. S. N. Y. S. Probably banded Both Both Both Band.d Handed Both ni-HHpiM'ured 11 of 34 purchased 5 of 34 purchaseer brood. 28 <-onU)ct^ ATTEMPTED REMEDIES 19 TABLE 2. RECORDS AND RESULTS OF GROUSE LIBERATIONS KNOWN TO THE INVESTIGATION (cont'd) Date Number liberated Source Wild or reared stock Where liberated By whom Banded or marked Results Nov. 1933 50 Alberla Wild Delaware Co.. N. Y. Robert Gerry Neither See below — 1936 by Gerry See below— 1936 by Gerry 1934 75 Alberta Wild Delaware Co.. N. Y. Robert Gerry Neither 1934 56 13 Alberta N. Y. S. Kesearch Center Wild Reared Franklin Co.. N. v. Conn, flill Robert Lehman N. Y. S. Neither Ituth Spring 1934 Only 3 recorded dead by Sept. Spring 1934 12 N. Y. S. Research Center Reared Hyde Park N. Y. S. BandfNl One contacted 6 miles away Aug. 1935 23 (.terry's Farm Reared Delaware Co.. N. Y. Robert Gerry Bundtnl •2 hunter killed. No mure seen Jan. 1936 23 Alberta Wild Delaware Co., N. Y. Robert Gerry 15 dead or killed. Rec- ord may include some from above. Identity baiM-d only on color Apr. 1938 12 N. Y. S. Kesearch Center Reared Conn. Hill N. Y. S. Both One found de^id. Several contacts Apr.-May 1938 84 N. Y. S. Research Center Reared l^harsalia N. Y. S. Both 17 found dead; 1 fe- male found with brood; 40 ctintacts. one 3 ^ Aug. 1939 17 Washinf^ton Reared Protection Is. Oregon Three- way Co-op. Unit NeithiT miles uwuy 1 br 1941 10 N. Y. S. Research Center Reared Ddniar IS. Y. S. Banded Some seen for several months in vicinity Winter 1941-42 223 of 381 106 3b Alberta Wisconsin Wild Wild Ohio Cons. Dept. Ohio Cons. Dept. R. FoUett Winter 1941-42 Winter 1941-42 Manitou Is. tWis.) Disappeared Spring 1942 40 Wisconsin Reared Neor Clare. Mich. R. FoUett 20 THE Hi F FED GROUSE l.\ THE MARCH OF TIME liberated by William Barnhard, a deputy game warden in Wisconsin, on Washington Island at the entrance to Green Bay in 1907-08. About 1911, 39 wild grouse were liberated on the Island of Anticosti in the Gulf of St. Lawrence, where previously they had been unknown. Al- though but 19 are said to have survived the first winter, they apparently bred and spread, for these birds are now fairly common over the Island*. While the demand always greatly exceeded the supply, no doubt many more birds were shipped, particularly during the 1920s. A record of those attempts, known to the Investigation, is given in table 2. Apparently, few permanent records were kept, for, if we may judge by present day experiences, scarcely a year goes by but that two or three such importations come to light. For instance, Robert Gerry liberated 50 grouse on his estate near Delhi, N. Y., in 1933. and 75 the following year, the birds coming from Alberta. Most recent records of importation are of 261 from Alberta, in 1940-42, by the Ohio Division of Conservation, for liberation in the southeastern part of that State. Not all the interest in grouse liberations is in the East. In August 1939 Arthur S. Einarsen, in charge of the Oregon Three- Way Cooperative Unit, liberated 17 hand-raised grouse on Pro- tection Island. Subsequent survival was carefully checked; one brood of five was seen the following fall. In 1942, however, there was apparently but a single survivor of the liberation resident on the Island. Another practice, the establishment of refuges, also became popular. These were organ- ized largely for bird protection in general rather than for grouse alone. Where fear was particularly strong that grouse might vanish entirely, such areas were selected primarily to preserve this species. The refuges served as breeding and nesting places so that grouse might be undisturbed by man. Occasionally the habitat was improved by planting open areas to food or cover-pro- ducing species. In some instances, also, the axe did its part in creating breaks in the forest cover. Thus opened up, the all-too-barren forest floor, in the Northeast, at least, usually gave way to a profusion of herbs, berries and second-growth sprouts. This proved to be a particu- larly attractive combination to birds, both old and young alike. Considered as protective oases in the vast expanse of huntable grouse cover, these refuges could have had only the most local of possible effects on the aliundance of the species as a whole. History records no startling examples of innnediate and satisfy ing increases following their establishment. Perhaps it is because man is, after all, but one of the major factors con- trolling the abundance of this great game bird. RISE IN IMPORTANCE AS A SPORTING BIRD No game bird in America has been more completely investigated than has the ruffed grouse. It was widely dispersed over the countrv. Its periods of abundance, followed by sudden disappearances, were disconcertingly mysterious. Its ability to maintain itself in close proximity to man and to nuikc him aware of its presence by its slightly unreal luil intrigu- ing drunnning. created for most countrv folk, living within its range, many pleasant ami mem- orable experiences. That it was sought after as food and was easily salable in the markets also contributed to llic iiitcrcsl which made these investigations possible. But it was the sportsmen to whom the skill and daring of the bird most appealed. Pur- sued by the occasional market hunter, it remained surprisingly tame. Once partridge hunting became popular however, the bird became wary and alert. It seldom proved possible to • Towntrnd, C R.. prr«onal Ii-tlfr In tin- aiillmrii. January 17. 1933. EARLY EFFORTS AT ARTIFICIAL PROPAGATION 21 shoot the last bird out of a covert, so adept did it become at dodging and screening its flight behind a convenient tree. Practice truly gave it more perfection until hunters came to have a very real respect for the bird and a genuine concern for its continued welfare. The men who had secured their living hunting partridges gave way to those who made it their recreation. Quick transportation made it possible for still larger numbers to enjoy the sport. The successful introduction of the ringnecked pheasant throughout portions of the grouse range, at a time when real sportsmen everywhere were particularly concerned about the ability of the rufled grouse to survive, caused some to shift their hunting to the newcomer. With each periodic recovery of the u])land favorite, however, devotees flocked back to it in ever in- creasing numbers. On the average these later hunters may not compare with the old grouse gunner in sheer deadliness, for they are neither such good shots nor do thev spend so much time in pursuit of the grouse, but they are among the best class of sportsmen afield. But what has really appealed to the sportsmen as potentially the most practical measure of all, were the attempts to raise grouse artificially. EARLY EFFORTS AT ARTIFICIAL PROPAGATION It is as true with grouse as with other game that, next to protection as a method of in- creasing abundance, man puts his faith in artificial propagation. Undisturbed, grouse are probably the tamest game birds we have. Harried sufficiently, they compare favorably with the wildest. But their nests, while hard to find, are not uncommon. Often they are uncovered by accident. Once found, the resistless human tendency to "see what will happen" has been the cause for literally hundreds of attempts to hatch out a clutch or two under a barnvard hen and to raise the resulting brood, (tables 3 and 1). These efforts began early. In a letter from John Bartram to George Edwards, dated July 15, 1750"°, that keen observer wrote, "Many have attempted to raise the young ones and to tame them, but to no purpose. Wlicn hatched under a hen thev escape into the woods after ihey are hatched, where they either find means to subsist or perish." While such empirical experiments were unquestionably repeated hundreds of times, it was not until 125 years later that the results found iheir way widely into print. In the fall of 1874, near Dansville. Maine. M. \^'. Clark tra|)ped four wild partridge, over- wintering them in a small cage. The following spring, he built an enclosure in the form of a circle, 10 feet in diameter, with walls five feet high, to protect the birds and prevent their escape. The lop was covered over with a conical tent. To make the grouse feel at home, he planted little pines and firs in close clumps, between which were scattered dry forest leaves. Even wild ruffed grouse may be tamed rather quickly in captivity and these proved no exception, for one soon made a nest in which were deposited 18 eggs. 22 THE RUFFED GROUSE IN THE MARCH OF TIME Clark tells us that the two females quarreled over the privilege of sitting on the eggs, a situation which more recent experimenters have seldom noted. Most likely both laid in the same nest. At all events, the eggs were subsequently set under two bantam hens, one ot which, unfortunately, was killed shortly by some predator. The other hatched four young grouse. Put in a small coop, she is reported to have developed a deep affection for them for which they made a poor return. Clark indicates they took little note of her anxious calls and that frequently at night he had to place one or two "lost" chicks under her. Almost prophet- ically, they all died, but Clark, little dismayed, carried on similar experiments for six long years before concluding that his failures were attrii)utable to a lack of proper food. TABLE 3. KNOWN ATTEMPTS TO REAR RUFFED GROUSE IN CAPTIVITY FROM EGGS COLLECTED IN THE WILD— 1879-1942 Year Expcrimcntator State EgRS set EgKs ha tehee Birds raised Year Expcrimentator State Eggs set hatched Birds raised 1879 Clark, M. W. Maine 10 ? p 1913 Job, H. K. Conn. 13.3 13 6 1880 ? 6 6 1914 McLean. Sen. 0. P. Conn. :t 6 6 1886 **l*enn" ? 12 ? 0 1915 Georpe. A. F. Graham, G. II. 30 27 ? 1890 0. O. S. 3 11 9 0 1914 Jeffries. G. A. N. Y. 7 6 5 1897 Newconie. \. ? 40 34 31 1915 ? p ? 190S Dr. C. F. Hodue Mass. 20 6 0 1916 Wolcott. Sen. Cunii. 7 7 4 190» 12 12 6 F. C. and 1905 p 15 0 1917 MacVicar, 1). p 3« 8 1906 12 10 0 1907 54 38 10 1916 Gerry. H. L. & NY. 13 12 10 1908 12 10 3 1917 Marshall. Wm. 11 p 8 1904 Manross, F. N. t.^oiin. 8 8 5 1928 Gerry, R. L. & N. Y. 10 9 0 1905 n ? ? 103(1 1933 Rae. r. 30 3 ^\ 0 3 1904 Rawson. C. L. .' ? ? 2 1934 1935 45 57 32 42 6 33 1905 Mass. n't. of Coils. Mass. 5 1906 Mprrill, A. 82 70 3 1919 Am Ciame Prot. A'n N. Y. 4.". 19 1 1907 56 50 4 1920 Allen, A. A. 37 2« 4 1 3 1931 42 16 0 1929 19.30 52 42 42 32 1 1 1912 Am. (jameProt. A'n Mass. 40 16 8 1931 1.38 125 57 1913 Torrey. II. A, 40 ? 11 1932 1933 78 56 63 51 58 17 1914 Mass. D't. of Cons. Mass. 136 97 18 1915 Torrey, II. A. 123 49 19* 1917 McVicnr. A. J. ? 80 31 1 1916 12 ? i> 1920 8 i) 1 1929 IlnuKhton. W. VI. N. Y. 13 13 0 1921 11 6 0 1930 14 II 9 1922 8 ? 0 1931 .36 ? 0 1924 12 0 0 1925 8 8 0 1929 Sines, II. J. Pa. II 1 1 9 1926 10 6 2 1930 40 25 0 1928 9 3 3 1929 11 11 0 1929 Turrill, I). Pa. 9 9 0 1930 13 13 8 1930 66 31 1 1933 20 15 U 1932 42 32 0 1934 20 19 11 19.30 N.Y.S. Cons. Oept. N. Y. 4 2 1936 MasK. D't. of Cons. Muss. 34 29 4 1931 Bump, (i. and J. 221 189 46 1937 Backus. K. K. 19 " 0 19.12 1933 Biim|>, G. tinil J., 365 681 285 (.12 111 126 1936 Mass. D't. of Cons. Mass. 51 30 2 1934 Holm, v.. 1031 892 158 1937 Sherman. L. B. 13 13 0 19.35 1936 Itnmp. (f. 701 371 601 .'148 127 47 1936 Maaa. D't. of Cons. Mass. 20 12 0 1937 Ilnlm. Iv. and 237 21 1 51 Torrey, J. A. 1938 19.39 Kordhain. ,S. 147 155 143 135 16 49 1936 Mass. D't. of Cons. Mass. 67 48 ? 1940 108 102 58 1937 Wrd Cons. D't. LeCompte, E. L. Md. 12 11 11 ? 0 1 1931 McCarthy, D. F. Pa. p ? 0 1934 Banta. L. Mass. i 3 1 1932 Keleher, C. M. Po. 30 29 18 1935 152 US 5 1933 29 28 3 1935 Hart. D. Mass. 10 10 8 1933 Ohio Cons. Dppl. Ohio 35 29 11 TrautiDan, M. !l. 1935 Mich. State Dept. of Cons. Mich. 9 7 0 1933 Conn. Statr ltd. of Conn. 16 It 0 Buhl. 11. D. Fisheries and (tame 1936 Trippensee, R. E. Mass. 28 26 1 Beck. T. 19.39 Follelt. R. Mich. 90 83 61 1933 McNamara. L. C. N.J. 22 16 11 1910 1941 313 110 81 ? 52 1933 Lces-McRae Coll. N. C. 50 50 10 ? Rogers. II. T. N. Y.A 1933 N.J. Bd. of Fish and Ciame Comm. Both. Kd. N.J. 15 15 10 Bendick. D. H. Brackett. Comm. Man.t Mass.t 1933 Nickerson. F.. Jr. Conn. 10 7 7 ? Batclle. J. B. OhioA 1933 Smith. R. G. Conn. 8 6 2 ? Kuscr. A. B. N. J.A ^Results of both wild and hand-reared eggs. ARaisnd some birds. fSonif r;uM<'(i ill large, open pens and sold. JSonif (•t.'^'s liiitched; none reared. §Oiu; liriiod started. ^Includes wild chicks rescued and raised. ^Week-old chicks caught in the wild. But then, as now, it was easier to try out the raising of grouse than to read of others' ex- periences— and infinitely more fun. "Penn"^ in 1886. "0. 0. S."°^ in 1891, and Commis- sioner Brackett"" in Massachusetts, about 189.5. all started grouse under bantam hens with- out bothering to leave more than passing records of their methods or their difficulties. Per- haps the most encouraging record was made by Newcomb. who, in 1897, collected 40 eggs, hatched 34 under hens and raised 31 chicks beyond four weeks of age. By the turn of the century it was generally established that grouse could be hatched from eggs gathered in the wild. Folks were not so sure, however, that the resulting birds could be raised to maturity, kept over winter and encouraged to produce fertile eggs the following spring. Long arguments around cracker barrels failed to settle the subject, but helped might- ily to make the job the more worth doing. It became a challenge to ability as well as an opportunity to contribute to science. With this in mind, Dr. Clifton F. Hodge"", head of the Department of Biology of Clark University, weighed carefully, in the fall of 1902, the problem of raising grouse on a large scale, and found it worth doing, "if merely to prove its practicability from the standpoint of science". So Massachusetts, long a pioneer in grouse rearing work, issued Dr. Hodge a permit to collect the necessary eggs from wild nests. A student, fired with the doctor's enthusiasm, placed the eggs in his hat and, putting it gingerly on his head, brought them to the Hodge home, there to be turned over to a broody bantam. This care was rewarded when six young partridges were hatched. None survived, however, beyond the first two weeks. Undiscouraged by the chorus of "I told you so" that arose, the spring of 1904 saw six birds raised from a clutch of 12 eggs. But, with these almost a year old and victory in his 24 THE RUFFED GROUSE /A THE MARCH OF TIME TABLK i. KNOVV\ ATTKMPTS TO HlvVU lU'FKED GROUSE IN CAPTIVITY FROM EGGS SECURED FROM HAND-RAISED BIRDS— 1875-1912 Year Experimentator State Eggs set hatched Birds raised Year Experimentator State Eggs set hatched Birds raised 1875 Clnrk. M. W. Maine 18 4 0 1940 823 412 224 IfiTS 16 6 0 1941 761 453 76 187'J 3 3 ,3 1942 684 350 113 1880 .3 6 6 1931 Coleman. W. B. Va. 39 21 4 1905 HilWSOII, (".. I,. p p 9 .5 1932 1933 30 12 16 4 8 0 1905 I^<«l^■.■, Dr. C. F. Mass. 10 5 0 1908 12 0 0 1931 1932 Turrill. D. Pa. llBt 61 61 34 0 3 1907 Mass, U<'i)t. (if Mas3. 21 8 0 1933 46 36 27 1909 ("oils. ~9 5 0 1935 ? 112 80 1910 Mnrrill. A. 21 ^ 0 1913 'I'orrey. II. k. 15 8 6 1932 Sibley. C. L. Coiiii. 14 ? ;> 1915 31 21 10 1933 p p 14 1916 68 31 1 1928 35 0 0 1933 Kelcher. CM. Pa. 49 6 6 1931 42 16 3 1934 20 5 5 1935 23 3 2 1936 12 3 0 1933 Ohio (>ons. Dept. Ohio 43 30 6 1937 Backus, E. E. 52 7 0 Trautman. M. 1!. 1937 Wood, F. W. 121 38 11 1938 159 30 6* 1933 Jaeckel, II. F.. N. Y. 140 85 6 1939 96 38» 15* Jr. 1940 28 0 0 1941 18 0 0 1933 Lees-McRae Col- lege N. C. 10 10 2 1915 Mclean, Sen. G. I>. Conn. 13* 12 ? & George. A. E. 1934 Ott. F. N. C. 10 10 2 1918 Wolcott. Sen. F. C. Conn. 31 27 .3 1934 Gerry. R. L.. N. Y. 19 0 0 1919 & MaeVicar, D. 72 48 2 & Rae, T. 1918 (Jerry. R. L. & Marshall, Wm. N. Y. 56 18 2 1935 Mart. D. Mass. 19 15 9 1937 Maryland ('ons. Md. 3 0 0 1927 Allen. A. A. N. v. 16 0 0 Dept. LeCompte. K. L. 1932 171 49 19 Md. 1933 265 96 42 JclTries. G. A. N. Y.i 1931 N.Y.S. Cons. Dept. N. Y. 25 2 1 1932 Hump. (■. & J. 165 97 75 Penusylv. Game Po.A 1933 Bump. G, & J. & 228 133 36 Comm. 1934 Holm. E. 825 258 37 Fredericks. W. 1935 1623 881 193 1936 Bump, G., Holm, 1385 .392 38 Houghton. W. M. N. Y.A 1937 E., & Fordliam, 1816 936 239 1938 S. 815 215 42 1940 Follctt. H. Mich. i> ? ? 1939 654 260 95 1942 260 ? .3 *Reault3 from eggs secured from the wild and from hand-raised birds. A Raised some grouse. tExact year unknown. JTwclve of the 20 birds producing these eggs were from wild-trapped stock. grasp, fate played the persistent doctor the first of a series of strange and discouraging tricks. A cat appropriated all Imt two for a midnight lunch although the grouse were carefully penned. Undaunted, the following year he collected a larger number of eggs. The two survivors of the previous year were, fortunately, a pair. One can imagine the excitement and satisfaction engendered when ten eggs were laid and five chicks emerged from them. Rv this time, too, cats had learned to give Pine Hill, where the Hodge family lived, a wide berth. This edu- cation was not an altogether painless process, however, for an assistant. Dr. Dellinger, was liaird before the Worcester police court, charged with cruelty to animals for having killed .-.^^,.'v-iv..-•"; *.•■••'--... ..— » • *— * • « .-^■^V tc-N^v ><\'''^=r=:iyi EARLY EFFORTS AT ARTIFICIAL PROPAGATION 25 one of the feline predators. The court held that the cat was a wild animal and therefore not entitled to protection by law except when treated in an inhumane manner. The doctor was released. Historians add that, thereafter, cats passed through Pine Hill, "only on the run". But not so with the indignant neighbors, some of whom, by now, were convinced that the only good grouse was a dead one. Feeling that the birds desired natural surroundings, a spruce tree on the Hodge lawn, 30 feet tall, was completely enclosed with wire and the seven survivors of the rearing season just passed were placed therein. With October's falling leaves came frosty nights, during one of which some acorns, filled with poison, were mysteriously tossed in the enclosure. After eating them, all the birds promptly died. The papers came out with articles about poison-fiends, but the cats were avenged and science suffered accord- ingly. Still determined and undeterred. Dr. Hodge continued his experiments for three more years. In keeping with all attemijts made before or after, success always lay "just around the corner". In 1907. his perseverance and good judgment in handling the experiments se- cured from the Carnegie Institute of Washington a grant of $,500 to further the work. That year, too, convinced that bantam hens were temperamentally unreliable and likely carriers of disease as well, he took a long step forward by raising a small group of birds under the primitive brooders of that day. The following year, grouse nests were extremely hard to find. Only one was collected. Though ten youngsters were hatched, seven were killed while feeding on striped bugs. From the adults of the preceding year, one hen laid 12 eggs, all of which proved to be infertile. Though he must have had some misgivings, as late as June, 1909, Dr. Hodge was still at- tempting to secure eggs from as wide a territory as possible "to learn to what extent geo- graphical conditions enter into the life of the bird". l\'o eggs were forthcoming, however, and he was forced, regretfully, to turn to other fields. It is significant to note that, at the end of seven years of careful experimentation, he felt fully equipped, both as regards experience and techniques, to breed and raise partridge suc- cessfully in ca|itivitv. Food, both for young and old bir.« iV -^-^ .J\>''o^ I ^^^>?^ ^C^vl 'fL':^ 26 THE RUFFED GROUSE IN THE MARCH OF TIME bantam: ])art in an infubatnr whose tendencies were characteristically unpredictable. Once hatched, these birds were then i)laced under the primitive edition of the now highly developed artificial brooder. The brooder promptly burned and so did the chicks. Some of those en- trusted to hens died of disease, diagnosed as blackhead, and others from a series of accidents which characteristically befall grouse in captivity. Only three survived the winter. By now, Hodge and Merrill were well known to each other. The former secured his ban- tam hens in Sutton, as well as a supply of food. Merrill, profiting by these contacts, placed each of his two hand-raised females on the ground in a separate pen, 8 x 60 feet. The cock grouse alternated between the pens. However, the birds proved adamant. One hen laid sev- eral fertile eggs but abandoned her nest when they were only partly incubated. The second hen laid 14, which were promptly removed to encourage her to lay a second clutch — but to no avail. Though most of these eggs were fertile, the chicks died during incubation or .shortly thereafter, as did most of the 50 which emerged from 56 wild-gathered eggs. Only four of these birds survived the winter. At Sutton, in 1908. a larger number of eggs than usual was inought in. perhaps 150 in all. However, details of the progress of the work are lacking. There is an indication of a heavy loss at the age of seven weeks. However, a fairly large number was still on hand at the beginning of winter, during which they were again reduced to four. These proved to be a cock and three hens, two of which were killed during the breeding season. The remaining one nested and laid five eggs which failed to hatch. Better luck followed some 20 wild eggs that were collected, in that 11 birds were raised to maturity. Once again, winter losses re- duced the flock to four. There is a period, sometimes a recurring one in the life of male grouse in captivity, dur- ing which they are extremely vicious. Apparently Merrill encountered this in 1910, for he states, "This year a cock was so vicious that it was unwise to mate in small pens, and a new experiment was tried. The four birds were wing-clipped and turned loose in a 2-acre enclo- sure. Two nests were found, one with 12 infertile eggs, the other with nine eggs, of which five were fertile and were hatched by a hen. Her last chick was lost by drowning when three weeks old." Then came one of those strokes of rare luck. Two wild nests were fomid and brought in. One was quickly lost, but. of the other, every one of the eggs hatched and the 13 chicks were grown in an artificial brooder to the age of six weeks without a single loss. Then, as though to prevent over-confidence on llic jiart of their keepers, all promptly died of an in- fection. A change of administration in Massachusetts terminated the work at Sutton. But other hands were ready to take it uj). The American (lame Protective and Propagation Associa- tion had just been formed and Harrv A. Torrey was engaged to take over the work of prop- agating grouse at their farm in South Carver. By 1912. they were readv and a start was made by locating four wild nests. From these, eight young birds reached iiialuiils in l')13. •\l)oiit the same nutnber of eggs was collected and iniough more birds were reared to make a total of 19 on hand at the end of that year. Here, significantly, occurs tiic first record of two grouse iuou;:iii to the sci-ond generation in ca])tivity. The birds were raised, for the most part, on the ground and with bantams in much the same manner as has been previously described. In spite of this early success, the farm was closed in January, 1914. and the 19 hand-reared grouse dropped from sight. Dr. George Vi'. Field, chairman of the Massachusetts Game Commission, was not at all EARLY EFFORTS AT ARTIFICIAL PROPAGATION 27 discouraged. Torrey, therefore, was placed in charge of the East Sandwich Game Farm and instructed to trap a wild brood stock of grouse. About a dozen were finally secured but did not breed well in captivity, so that the main dependence was again on wild eggs. Dur- ing this and the succeeding year, at least 150 of these were obtained from one source or an- other, mostly by the game wardens. Sensing a long and difficult time ahead, however, the "powers that be" overruled Dr. Field and officially terminated the experiments in the spring of 1916. Without so much as a band being placed on them to help determine future survival, the birds raised that summer were liberated. However, Torrey's heart was with the grouse to the last. Thereafter he never lost a chance to pick up a clutch or two of eggs and to rear as many as possible of the chicks, usually with bantams. In 1933, nine of the birds, which Torrey raised personally, were graciously contributed to swell the propagation experiments already under>vay by this Investigation. The following year Charles Dimniick. long-time grouse enthusiast, returned from a fishing trip in Vermont with two clutches of eggs. All in all, he carried these well over 300 miles. At his request Torrey set these under two broody bantams, one nest in either side of an orange crate. Eleven of the birds that hatched grew to maturity, cared for by their foster mothers, only to fall prey to great horned owls which decapitated them while they vainly tried to fly through the wire of their pen. A decade earlier. Dr. Arthur A. Allen had demon- strated conclusively that poultry could usually be depended upon to carry diseases fatal to grouse. That these birds were raised at all, therefore, was a tribute indeed to the care and perseverance with which Torrey carried on his work. Two others among a host of experimentors stand out during this early |)eriod — John Burnham. long-time president of the American Game Protective Association, and U. S. Sen- ator Frederic C. Walcott of Connecticut, one of its most enthusiastic backers. Roth were dis- satisfied with the cessation of Torrey's South Carver experiments. To each came the oppor- tunity to further the cause. With the establishmenl of the U. S. Biological Survey's experimental fur farm in northern New York, George Jeffries"" was appointed its operating head in 1915. Backed by Burn- ham's enthusiasm, Jeffries secured a few clutches, in 1915, from which he over-wintered six or seven birds. This experiment was repeated the second year, with like success. Though, at the time, few folk worried about ground-borne diseases, the fact that Jeffries selected land on which no poultry had run for over 20 years, unquestionably was one secret of his suc- cess. The World War. in 1917. removed Jeffries to other fields and terminated the experi- ment. Meanwhile, in western Connecticut, on the Childs and Walcott preserve, a Scotch game- -■'^-""■'"■"''■■■'' " 5*- :--0}»ft^-., ■■•>v'.''' 28 THE RUFFED GROUSE L\ THE MARCH OF TIME keeper of the old school. Donald MarVirar"". under the a|)|)raisinn: eye of Walcott. aban- doned small coops and enclosures in favor of wide ranges to raise four grouse out of seven hatched in 1916. The foster mother, a game bantam, was wild and excitable, and the birds required constant watching. It is interesting to find that a chiiunnrik is credited with killing two youngsters, three weeks old. In addition to the insects and. later, the seeds and berries which the chicks picked up. the usual feed of hard-boiled egg yolks and fine game meal or cracker crumbs was provided. Given their freedom, one bird was killed in the kennels and the other three disappeared early in September at the beginning of the '"crazy flight" period. Only one ever returned. MacVicar's whole emphasis seems to have been on the maintenance of natural conditions. The following vear. three clutches of eggs were placed under bantams in coops .SO yards apart. The hillside was dotted with evergreens, birches, ferns, wild grasses and huckleberry bushes — an ideal natural grouse habitat. Because the weather was wet and insects not too |)lentiful. ant eggs and cottage cheese were added to the diet of the broods. As with [)heasants, when a couple of days old. the birds were liberated from the coops in which the foster inolher was still confined. All went well, when difficulties, characterized by diarrhea, resulted in the death of most of the birds. MacVicar attributed this to stale food left in the vicinity by a new assistant, though later experimenters have encountered the same trouble under a variety of conditions. The eight birds that pulled through were placed in a special pen covering about three- quarters of an acre. One bird escajied and another was killed by flying into the wire. Of the six birds left, one of the three males began strutting in Scjjtember. So savagely did he dominate the enclosure that the other two males had to be fed separately and eventually were killed by him. The three hens nested and laid 31 eggs, from which 27 chicks were hatched. Thus again were the polygamous and often domineering tendencies of the bird, in captivity, demonstrated. There are many others who might also be mentioned, particularly Manross of Connecticut, Herbert K. Job. and Harry Rogers, builder of New York's pioneer Sherburne State Game Farm. All of the exijerimenls, however, followed more or less the same pattern. Foster mothers or primitive incubation and brooding equipment were used, the eggs being largely collected from wild nests, and the birds, when hatched, being allowed to run on the ground in small enclosures containing more or less natural cover. Disease was only sporadically rec- ognized as a cause of loss, particularly among the young chicks, and then it was traced back to its probable source, the hen. Emphasis was placed largely upon providing the birds with as nearly natural surroundings as possible and. particularly, with natural foods. Eggs were usually removed from the nests when incubation was already well along and, therefore, most of them hatched well. Usually the birds started to die almost immediately, however, and the losses continued at a heavy rate until they were three or four weeks old. .Vfter that, a few birds might lie lost from disease or the whole gi(iM|> wiped mil bv an epizootic. So primitive were the conditions under which the grouse were raised and so consinning of lime and atten- tion were the teihni(|iies required, that most experimentors ascribed their losses to prevent- able causes of one kind or another, thus encouraging further trials. Now, however, the |)i(tiire changes. Poullrv science was making rapid strides, not only in perfecting incubation and brooding equipment, but also in the field of nutrition. Pathologists, likewise, learned how to recog- nize and prevent many serious avian diseases. The stage was set for someone to lift these EARLY EFFORTS AT ARTIFICIAL PROPAGATION 29 empirical experiments out of the realm of chance and to make of them a careful series of planned projects. That man was Dr. A. A. Allen, Professor of Ornithology at Cornell. Emphasis in the conservation picture was rapidly shifting from protection to the raising and liberation of game birds as an assurance of better shooting days ahead — the result largely of unprecedented successes in rearing and stocking pheasants. The American Game Protective Association attempted to further this movement through establishing an experimental game farm at Cornell University with Dr. A. A. Allen in charge of the research program. It was Dr. Allen's thought to contribute to the success of this venture by developing a technique for the artificial propagation of ruffed grouse. DR. A. A. ALLEN. DEAN OF AMERICAN GROUSE BREEDERS, WITH SOME OF HIS HAND-RAISED BIRDS When the farm was taken over two years later by New York State as a pheasant propaga- tion unit, Allen, "in full ignorance of the real problem, spurred on by the partial success of others ..." continued for a full 13 years a series of most significant and productive ex- periments. The bulk of these are well described in a paper entitled "Ten Years Experiments in the Rearing of Ruffed Grouse in Captivity"" presented before the Sixteenth American Game Conference in 1929. It was he who first successfully controlled many of the diseases, which had plagued earlier breeders, by rearing and holding his birds on wire. Likewise, his experiments led to the development of feeding formulas which furnished a sound basis for propagation experiments subsequently carried on by the Investigation. The colony breeding 30 THE RUFFED GROLSE I\ THE MARCH OF TIME en Z. o < O H CO V > z a O e o D f.s II u 1901-1908 1904 1904-1905 I905-I910 5 ooo CO- — c"» £ L- u-: r- r; — 4 ^ a: 3 — ci ei rj — — CI CI -^ -. c- c^ CJCI C 0^ "'2£ ? c- d- -ICICJ CJ -1 CI 5 50X Factors stiidird in some detail )uauiij£euBi^ XX::: :X sapXo : XX::: :x :x uoi^nSBdojd leiagrjjv X ; X X X XXX ■X :X X X : X : now X X X X : X :X : : X X saesasiQ X X X XX X xxxx : X sannaog X X X X : X X :x X J3q,B3M| X X X X X : XX XX: X X X-ijOBdea aApanpojddu X X X X : X XX X Bjiqeq isjaaao X X X XX XX X XX X X X jajpqS X X X X : :x XX: X X POOJ X X X XX XX X XX K X X jCSoioieXq^j : : X : : 2S. 1 1 55 5 5 5 ^1 -:,i 1 h III J 1 1 III |=llll |i| 1 ills 1 i ii III 1 M 4tt it^iti I'^t I s^sit d| ys SSu S 2 S Szz zz zzz z z z S^SS^ zeS zS«i In charge of. or reported by 1 1 ill 1 1 1 ill li d 1 j Ji 4iiij.il! ill uilz J en < &- : :S : -s : :S : il :E :^ • a :•! :£ '■£. :S ■S : i = > z c ; C < : 1 5 d £ \ ^ ! i fill ■ m ■ .*iJ r £ C 'C •< . . . > . 1 ill ; 1 • ^^ ■ < :<| : « ■ a ,S : > ^>c . 'fe'^l '■ :i ^^3 : i-U~ : ill y JZ'J" Jl •s : : : : J : : : : tSf'S • • • 4|:|| i> :-! <-= :i3 I ■- 0 = = ■ "i ■ s dO .2 = 31 II n jj e 'K > ■5 3 c 0 Hi J a :- niu SszzS i 1 s 1 ■5 1 HlimuiaaUc; I :X :x XX x xxx :X : Xu!:|3(r|K>i| j : spjoaou 1 : X : OJinuiioiiNaii^) j > t : X :x X Xjo]njaqir] | ; fiuotlDAJintqo pi^lj] 1 :xxx XX X X : X XX :x )U0UldO|UA9Q 1 : uortnUedojd iniogivv IX ■y.x > < X x* ( : s> « : :x X 1 EARLY EFFORTS AT ARTIFICIAL PROPAGATION 31 la If 2.S II 1935-1936 1935-1938 1936-1937 1936 1936* 1936 1938-1939 1938 1939 1939-1941 1939-194(1 1939-1942 1940-1942 2 2 o c c a- « « CI •* h P )uaui3:fT!ue|^ >^ X ;x :X X sa|3Xo X X :X uoiieSodojd lejjyiijy neiv :X : X XX :X X sdsuasiQ :XX :X X X II uainiaua | XXX X X jaqiB8A\ 1 : :X X X X^pcdea aAj^anpojday j X ;X XX X :x X X S)!quq [Bjauaf) | ;X X XXX X X jaiiaqsi : : : X XXX X iJooj 1 X :X X : X XXX X X XXX xaoioiB;«qj 1 : :x X : : : : : : x ^8 =' J 0 S S t 3a.4^^«.c»a^ ffl 1 i 1 £ ^ - si tc a^ * 5 S =i-c P _ ^ 3 1^ i 1 1 > 1 E- — : ■ L ■j Z ! c ■j > 1 a ■■u Hi 385 E- H 1 .5 "Ei_ * m" C c" d 'la 0'- : : g -1 1 i j ? s ? - ^£ « ^ i ce S "S ! -■ 2 £ ! 1^1 = s. •= «■ = 1 ^ ^: 1 — oi ai : )Z 3 a iS -ll- fill = !*« = ,«Z32 ! f ! . c " c :i ■ >> 3 *0 a >> suamioads 1 : : :X : :X : : : : :XX : X XX : x || nuinDcnsau 1 :::::::::: :X : : spjooay 1 ::::::::::::: : : : ajiBuiiopsan() | ;;:;::: :X :::: : : : : xjoiBjoqai 1 :::::: :X ::::: : : :>< suoiiBAjasqo pjaij | X(oul 17 iiiclio in Icnglli anil ha\c a wing-s])rcad of sotuc 23 inches, (p. 98). Grouse feeding habits are largely neutral from an economic standpoint, but their budding of fruit trees has, in some sections, been serious enough to result in bounties in at least two states, (p. 102). As a game species the grouse is an ini|)(irtant economic asset, both financially and recrea- tionally. (p. 103). Inherently endowed with a ■"bnmp of curiosity", the human animal has been interested in other forms of life about him simc long before the dawn of history. As one expression of this interest. primiti\e man drew pictures of familiar species, or carved images of them, to decorate his caves, lodges and implements. Crude at first, his skill constantly developed and, with it. a desire for more than mere reproduction of the form of the creatures he encountered. Consequently, students have care- fully described their appearance and probed their anatomy, plotted their occurrence over the 38 THE SPECIES ITS TAXONOMY, RANGE. BIOLOGY. & ECOXOMIC IMPORTANCE earth's surface and pondered their evolutionary relationships. More recently, preliminary excursions have been made into the vast fields of animal behavior and experimental physiology as they apply to wildlife species. Unknown to the white man until after the discovery of North America, the ruffed grouse has since received considerable attention, especially because of its economic importance. The resultant information has, however, been widely scattered in the literature. It is the purpose of this chapter, therefore, to bring together the more important of these data as well as what new material has been brought to light by the present Investigation. A logical starting point is the bird's taxonomy. TAXONOMY A major step in the development of man's knowledge of the universe has been the cata- loguing of other forms of life. Largely haphazard at first, tliere has gradually evolved a definite system for classifying any animal or plant according to its evolutionary relationships to other species. While anatomical features are the fundamental basis for such classification, a descri])tion of the external appearance of a species is a basic preliminary, and is also important to the general wildlife student. Description The ruffed grouse is a brown, fowl-like, woodland bird about the size of a bantam lirii. possessing a dark, partially concealed ruff on each side of the neck and a faiishaiied tail having a broad, dark subterminal band. Although referring primarily to the type form Boiiasa iiiiibclliis iiiiihclliis i Linnaeus I *. the following detailed accounf^ applies in general to the species as a whole, the differences between subspecies being largely a matter of degree. ' "Adult male: Top of head with many narrow bars or spots of rusty brown, black and sometimes whitish; rest of upper plumage mottled and variegated rusty brown and whitish or buffy. with some black marks: inner seciiiularics and wing-coverts commoidy edged or tipped buffy-whitish, often spotlcil with black: primaries dark brownish-gray, marked and spotted with buffy or whitish on narrow outer webs: back and rump willi numerous oblong or lance-shaped pale, black-edged shaft-spots; neck ruffs usually black with green and purplish reflections, in some cases reddish-brown; tail bright reddish- brown to gray, barred regularly with from 6 to 11 narrow blackish bars and one broad subterminal black bar: chin and throat buff, loucr and lateral feathers often tipped dusky; wing linings and axillars brownish-gra\ and while; rest of lower plumage whit- ish, grayish-white or buffv-white. usually tinged buffy on lower neck and breast, with many bars of deep buff, brown or black, all darkest on sides and flanks, often fading and obscure on breast and belly; buff and white j)redominate on under tail-coverts; bill dark brown; iris hazel; feet dark horn. "Adult jnitale: Similar but duller, with somewhat lesser ruffs and shorter tail: iris, bill and feet as in male." » A.O.U. Check Li.l No. 30tf>'. A while this acroimt ifi qiiotrd from Furliiit.li'*'* exii-llrnt description* hove nUo been |>iibliithe(l by Roberls*"", Chnpmnn"*. and othcm. t Sec immmnry o( siibiiprcirir iliffercnces. p. '18. IJ^'^^X TAXONOMY 39 How TO Recognize Sex* How to tell male from female has always been a matter of interest among sportsmen. From the stag and wild boar of the mediaeval sons of Diana to the grouse and pheasant of contem- porary scattergun enthusiasts, there has been a tendency to prize male specimens more highly. But, while the males of many game species have distinctive adornments or plumage, in others Nature has been much less lavish. Among the latter is the ruffed grouse. Positive determination of sex must, of course, depend on examination of the gonads^. In birds, this is often difficult, especially for the untrained observer, since the organs are internal and very small, except during the breeding season. Therefore, a knowledge of any features of coloration, external anatomy or actions, which tend to distinguish the sexes, is desirable. The Investigation has had unusual opportunities for observing such differences in grouse, particularly with respect to artificially propagated stock at the Research Center. Constant association with birds at this unit, as well as in the field, has led to the recognition of certain characteristics as indicative of male or female '. Activities such as drumming and nesting are, of course, quite definite criteria of sex. but features of feather marking and bodily propor- tions are much less distinct. The latter will be considered first. Before discussing these differences in appearance, it should be pointed out that no one characteri.stic, or even a combination, is infallible. In general, they represent extremes of variations which exhibit all degrees of intergradation. Thus it would be a very unusual bird which would |)()ssess all of them as described for its sex, yet most specimens will have a majority. One must also realize that the characters are largely relative and, unless one has a suitable basis for comparison (either a series of other specimens or wide personal experience), judgment is diflficult. Furthermore, the features associated with bodily proportions apply primarily to living grouse, although many of them can, with care, be used for freshly killed birds. The accompanying illustrations will be helpful in visualizing most of the differences dis- cussed. In studying both paintings'' and marginal sketches, one should keep in mind that the extremes have been pictured in order to emphasize the various characteristics. The First Things to Look For Perhaps the most diagnostic external feature is the tail. Birds having the broad, dark, subterminal band uniform over all tail feathers are almost invariably cocks, only one excep- tion having come to the authors' attention. But, if this band is broken on the two middle feathers, the bird may be either male or female. In such cases, the length of the tail should be determined. In males, this appendage tends to be over six inches long: in females, it is shorter. Exceptions are not unusual, however, and a more applicable rule would be to con- sider only those of six and a quarter inches or over as male, and five and three-quarters inches or less as female. When the tail band is broken, the mottled design in this area is apt to be coarser among the cocks. Moreover, a closeup view of the tail will show a tendency in * This account has hrm written hy Fred. Everett, who. while studying tlie birds at the Resrarrli Center and painting the colored illustrations for the book, discovered a number of previously unrecognized differences. A See discussion of Anatomy, p. 59. _ _ t The material upon which this discussion is based was derived from the stock inhabiting New York, which involves only the subspecies itmhetlus and logato. It seems probable, however, that the characters noted will be found to hold, in general, for the other races. t See color plate facing p. 40. 3 O a o a P o [d u ni>l found in the arid Harney Basin of central Oregon, the interior of Washington, or the Snake River Plains of southern Idaho. * Taken largely (r have always been sparse. Many of the early exiiedilioiis to ihis region failed to record a single specimen. Yet they apparently occurred in small lunnliers over a (■(>iisiderai)le area. The general situation seems to have undergone scant change, luil in Colorado no records' since that of Sclater*" in 1899 have come to light and it appears th;it the species is no longer • Crnniip wt-r.- f»iKfT««fiiIlv inlrn.lucrd nn thi* Itlond in 1911. .... i i .. A K.lim.tr.I I.V nrnnilt. R.. l'r..f. Zoology, Univcr.ily ol Mi..,niri. at nol .,v,T 100 cl.i.iv,. o( rrcciil Lbrmlion.. ,.T...n»l Irtlrr to !he authors, Jonunry 23, 19.t2, t Vpattpr, R. K.. nrrsonal Ir-ttfr tn llii* uiitlmm. Jjiniinrv !.'>. 1912. t IPailFT, l\. r,,. JPi-imiliui ii-ii.-i 1 "". "• J "- .■ - -- t Walcrfirlil, S. W.. iicrnoiml lrtt.T to tlir oiithnrH. .\pril 2, 1942. } Hcrjtold"', in 1928, nolo it m a "rare resident" but li«t« no specific observations. RANGE 51 o Boundory of Former Ronge Records Since 1925 Records 1900" 1925 FIGURE 2. DISTRIBUTION OF RECENT RECORDS OF RUFFED GROUSE OVER ITS FORMER RANGE IN THE NORTH-CENTRAL AND MID-WESTERN STATES present. In the East, various of man's land use operations have forced the birds to retreat somewhat to the higher mountains in Virginia and the southern Appalachians. In the North there is no reason to believe its distribution has undergone any important change. Since it is a species of sub-climax associations and is not intolerant of man's activities, so long as suit- 52 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE able cover remains, it is probable that its present range will remain essentially stable for some time to come. Of interest is the inclusion of two states, Nevada and Oklahoma, in which the presence of grouse has hitherto been unrecognized. With respect to the former, although substantiating specimens are not at hand. Rassmussen has recently reported* the scattered occurrence of the species in the White Pine Mountains, the Ruby Mountains and another small area in the northern part of the state. For Oklahoma, no records are available, but the former existence of the bird in the Ouachita Mountains to the western limits of Arkansas clearly indicates that it must have occurred at one time along the eastern margin of the state. In compiling the data upon which figures 1 and 2 are based every effort has been exerted to assemble the available material on the subject. In addition, the game departments, as well as other recognized authorities of the 52 states, provinces and territories in which the species at one time or another has been reported were canvassed by letter during the winter of 1941- 42. The replies received have been invaluable in supplementing published accounts. Neverthe- less, errors are inevitable where definite records are scarce or lacking and one must rely on judgment. This is particular!) true in northern Canada and much of the Rocky Mountain area. The authors believe, however, that such discrepancies will prove to be minor. Beyond this, certain statements which warrant comment are to be found in the literature. In 1831, Audubon"' indicated the occurrence of grouse south of Tennessee along the Missis- sippi, stating "but as you approach the city of Natchez they disappear". This seems to have been the basis for the inclusion of Mississippi in the range of the bird as given by Baird, Brewer and Ridgway". No specific records were given by either author and, since subsequent confirmation is lacking, this section has not been included. At the same time Audubon also ascribed the species to Texas but gave no particulars. Here too there is considerable doubt. The only other reference to its occurrence in this state is a note in Forest and Stream for 1878"° concerning its reported presence in the "Brown Sand Hills almost in the center of the Staked Plains west of Cedar Lake". It seems quite probable that this was a case of mis- taken identity since Cooke"^ states definitely that the species did not reach this state. On the west coast, sabini was stated by Cooper'" to occur as far south as N. lat. 34° 30', and northward, Macoun'" credited it to the entire coast of British Columbia as well as the Queen Charlotte Islands. Since these records now appear to have been based on misinformation, they are not included. One possible source of error was a specimen of this subspecies col- lected by Bishoff and cited by Nelson""" which was labeled Sitka, Alaska, but which, it has since been determined"*, was undoubtedly taken elsewhere. Another aspect of this subject is the sporadic occurrence of lone individuals outside the normal range of the species. In the North, as discussed elsewhere^, there appears to be a considerable area where stragglers may not be unconnnon although observers are so few that data accumulate slowly. Thus Ross™ recorded the species about 1860, at La Pierre House near the mouth of the Peel River in the Mackenzie Delta region where it probably wandered from the valley of the Porcupine. In 1878. Bell'' reported it from York Factory at the mouth of the Nelson River on Hudson Bay. More recently, Clarke'" took a specimen at Fort Reliance at the eastern end of Great Sine Lake where natives regarded it as quite rare. Similarly, Wayne*" cites a record for Camden. South Carolina in 1901. It is possible. too, that Audubon's experience in Mississippi may ha\c inxolved observations of this kind. * RaimuMrn. D, I., iifrnonal \ft\CT In the aiilhor*. May 22, 1942. A Srr ilitruftnion undrr Ditlribuliuii and Rrlmive Abundanro. p. 56. DISTRIBUTION AND RELATIVE ABUNDANCE 53 As stated previously, it is not within the scope of this report to undertake a detailed dis- cussion of subspecific relationships. Therefore, only a very generalized picture is presented. For each race, the type locality* is marked in figure 1. It is probable that the area of former range throughout the Mississippi watershed, east of the Great Plains was largely occupied by umbellus, while in Colorado, umbelloides was the predominant form. An important aspect of the distribution of these races is that between no two does a clear cut line of demarcation exist. Rather, one finds zones of intergradation where individuals of one subspecies occur side by side with those of another and where birds of intermediate coloration are also common. The subspecies involved in such zones of intergradation are. in general, those whose basic ranges are adjacent. In the case of togata, however, certain interesting variations exist. In the East, it invades the area occupied by umbellus in the form of a long arm reaching down the Appalachian Mountains where Transition or Canadian conditions occur. In the West, it ex- tends along the eastern margin of the Rockies through Montana and Wyoming, and in British Columbia, it crosses the mountains, inlergrading with sabini in the lower Fraser Valley and with umbelloides northward, at least to the Cariboo district'". This suggests the possibility that, from the standpoint of evolution, togata may represent the original race while the others are more recent modifications. DISTRIBUTION AND RELATI\ K ABUNDANCE^ The ability to occupy such an extensive range illustrates a highly important element in grouse ecology, namely, that the species is typical of sub-climax forest vegetation — shrubs, second growth and forest edges. Accordingly, it finds suitable conditions associated with the developmental stages of several climax types (biomes) rather than being restricted by the limits of any particular one^. From Alaska and northern Canada, where it frequents the spruce timber side by side with the spruce grouse, it seems equally at home in the redwood forests of the Pacific Coast, the aspen groves of the prairie country, and the mixed woods of the Northeast. Furthermore, early records indicate considerable populations to have inhabited the hardwood swamps of Indiana and Ohio, and Audubon"^ even states he encountered these birds in the canebrakes of the Mississippi. At the same time its distribution within this teriitoiy is far from uniform. Such a range of necessity includes a host of local areas unsuitable to grouse but impossible to indicate on a small scale map such as figure 1. Furlhermorc. the lialiitats in which the bird is found run the gamut from good to poor, resulting in wide variations in relative abundance irrespective of fluctuations associated with cyclic behavior. The following discussion of these features is divided into two parts dealing with the range of the species in general and with New York State respectively. Entire Range Conditions approaching the optimum for grouse* are found mainly across southern Canada, northern Minnesota, northern Wisconsin, northern Michigan and south through New England. New York, northern Pennsylvania and portions of the Alleganies I figure 31. Even here, however, are many areas unfavorable to grouse. British Columbia and western Alberta, in particular, present a complex interspersion of timbered valleys and alpine peaks — the one supporting grouse, the other not. Along the edge of the prairies in southern Alberta and * See discussion under Taxonomy, p. 47. A Bv Robert W. Darrow. t See also Pilelka"". t Sec Chapter HI. p. 110, and IV, p. 229. 54 THE SPECIES^ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE Saskatchewan (and furiiierly over much of the area now within the states of Ohio, Indiana. Illinois and Kentucky, which would have been included in the region under primitive con- ditions) tracts of treeless grassland are found in the midst of good grouse cover and vice versa. Many other minor exceptions also exist, such as the sand dunes of Michigan and the alpine mountain tops of the Adirondacks and \^liite Mountains. Grouse have, in general, always been most plentiful within this portion of their range, but. though the average is relatively high, some sections are more productive than others. Thus, well watered areas are more productive than dry; moderate slopes are preferred to steep; and cover containing an apjjreciable admixture of evergreens is better than pure hardwood. Areas intensively occupied by man support relatively fewer grouse than those where the habitat has been less altered. Beyond this grouse find their needs most adequately supplied in coverts comprising an irregular pattern of tvpes and having a more or less open crown but considerable under- growth. They prosper best in habitats associated with sub-climax forest types. The greatest primitive populations were probably produced along the water courses of the region of mixed coniferous and deciduous forest, about the swamps in the hardwood sections, and in portions of the prairie border of the middle-west and the southern prairie provinces. Burns and wind- slashes, however, also broke up the uniformity of the cover and produced highly favorable conditions. Conversely, extensive tracts of uniform type seldom have been conducive to high densities of population. Thus grouse abundance increased materially in the second growth following settlement and lumbering, although, of course, they disappeared where land clear- ing was intensive. Some of the most highly productive coverts today are found among the hill farms of New England and southern New York, especially where many of the clearings are beginning to revert to overgrown-land as a result of abandonment. Elsewhere grouse populations are more sparse and. in many sections, the distribution of favorable cover more scattered. Through its southward extension in the Appalachian system, its numbers are relatively low. While thev formerly may have been more abundant locally, especially where Canadian conditions existed, the effects of lumbering, burning, land clearing and grazing, through eliminating the evergreens, destroying the undergrowth, and drying u|i the streams, have resulted in their now being relatively scarce. A recent rejiort* from Ken- lucky, however, states that grouse are re-occupving manv areas following better coiitrdl of burning. Throughout tiicir range in the western states, grouse iiave never been luinuMous. Reports indicate that, on the Pacific Coast, they are most plentiful on Vancouver Island. The distri- bution of the scattered birds in Utah and Nevada is verv irregular and changes cdtiliiuialK although the general extent of their range remains the same. Here they are found cliicfiv where aspen and willow groves occur in canyon bottoms at moderate elevations. In the southern Rocky Mountain region the area occupied bv grouse lies as a band along the moun- tain slopes, its lower level being roughly defined by the occurrence of evergreen timber, its upper limits falling somewhat below timberline. In its former range in Colorado, this band lay roughly between 6000 feet and oOOO feet. To the northward it gradually lowers until, in the northern states and Canada, the valleys comjjrise the grouse range and more and more of the mountains project above timber-line. Throughout this region, especially in northern British Colum])ia. Yukon Territorv and Alaska, local areas unsuited to grouse (indicated onl\ approximately on tiie map) are frequent. To tiie north, alpine conditions arc chiefly rcsi)on- sible; to the south, arid intrusions of the Sonoran Zone in the river valleys are also involved. ♦ Wolcorielil, S. W.. pcrnonal Irltrr tn the autlinrfl. April 2. 1942. FIGURE 3. RELATIVE ABUNDANCE OF THE RUFFED GROUSE THROUGHOUT ITS PRESENT RANGE 56 THE SPECIES— ITS TAXO\OMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE Along the northern border of its range in weslern Canada, the forest cover is found pri- marily close to the margins of the streams and lakes, reaching like tentacles into the tundra and leaving expanses of nearly treeless moor between. South of the area between York Fac- tory at the mouth of the Nelson River and the east end of Athabasca Lake, grouse are regu- larly found in small numbers along these projections. Hut northward, the relationsbij) between grouse range and the tree-line diminishes rapidi\. In this section, tiic line shown on the maps represents the boundary of the area within which grouse are fairly constant in their occur- rence, although their numbers are low and many local areas, from which they are absent, exist. Nevertheless, grouse occur sporadically beyond this boundary, often to considerable distances. Clarke* even suggests that the Mackenzie Mountains should be included in the area throughout which stragglers may be found. This region has been so little explored orni- thologically, however, that definite statements are impossible. Especially noteworthy records are those at Fort Reliance at the eastern end of Great Slave Lake, and La Pierre House near the mouth of the Peel River. A similar situation, although the zone of termination is probably narrower, exists along the southern edge of the great plateau of Labrador and northern Quebec. Here, however, the extensions of suitable habitat project up the streams where they have cut valleys in the southern coastal escarpment, rather than downstream as in the case of the rivers flowing northward into the tundra. On this plateau there appears to be even less relationship between the extent of grouse distribution and the limit of treegrowth. The reason for this is that the tree line'"" is considered to include a vast area supporting only a scattered and stunted growth of black spruce, balsam fir, birch and jack pine, interspersed with tamarack swamps and large expanses of bare Precambrian rock. Nevertheless, most of the region is still unexplored ornithologically and our concept of grouse distribution in it may have to be considerably revised as further data are secured. But below this northern border zone lies a broad belt stretching from Labrador to the Cana- dian Rockies and into the Yukon valley where conditions are such that grouse may occur most anywhere, but where, although generally distributed, they seldom become numerous. New York State With the exception of certain Adirondack mountain to|(s which rise to timber-line or above, and the coastal marshes of Long Island, the entire area of this State originally constituted suitable ruffed grouse habitat, varying, of course, in productivity. Today, after three cen- turies of settlement, one may still encounter this species in every countv. with the exception of the Bronx, New York, Kings, Queens and Richmond, all of which lie williin the area of New York City. In fact, it is probable that densities of population now attained in many localities considerably surpass those of primitive times. Although the species often manages to persist in small nuinliers under marginal condi- tions, it naturally thrives best where it finds an abundance of its essential needs. In this category, in addition to an adequate area of forest or woodland cover, are an admixture of evergreens and suffipen(lii, p. 721. t See dinnifliiion of Clamiiriration, p. 16. t See dtncinininn of Voice, p. 98, PTERYLOGRAPHY 59 and hard-coated seeds. Branching from the oesophagus, between the mouth and the gizzard, is the crop which serves chiefly as a storage pouch for freshly eaten material. At the junction of the small and large intestines are attached two blind ducts or caeca several inches in length. As with birds in general, the two testes of the male are oval bodies lying on either side of the backbone and against the forward portion of the kidneys on their lower surface. During the breeding season these organs become greatly enlarged but at other times of the year may be so small as to be difficult for an untrained observer to identify. In the female, the ovary, resembling a bunch of grapes, occupies the same position as the testis. Normally, however, only the left ovary develops in birds. It, too, becomes much enlarged during the breeding season. Abnormalities of the sex organs are not infrequent. Thus, under certain conditions, the ovary may wither on the left and the rudimentary organ on the right develop into a testis. The effect on the plumage and other secondary sex characters is often remarkable. PTERYLOGRAPHY* The presence of feathers is one of the major distinctions of birds as a group. Among the various species of the earth, wide modifications from the primitive type have developed. Such differences, chiefly involving arrangement and types present, are used extensively in studying evolutionary relationships. In the Appendix'^ is included a detailed account of the pterylography of the ruffed grouse, of which the following is a brief resume. Although protecting practically the entire body, the feathers of a grouse, like those of most other birds, grow in well defined areas called tracts rather than indiscriminately over the sur- face. These tracts are bilaterally symmetrical and in them the feathers are usually arranged in two series of rows, one at an angle to the other. This pattern can be easily seen in a plucked bird. Tlie intervening areas of bare skin are termed spaces and are covered by feathers situated in adjacent tracts. The development of such spaces in birds, as well as the restriction of the feathers in tracts, undoubtedly facilitates better bodily efficiency through allowing great- er concentration of blood vessels. The brood spot is principally a space although some contour feathers are plucked by the hen along its outer edges. Of the many types of feathers found among birds, the grouse possesses neossoptiles. mesop- tiles, teleoptiles, and filoplumes. Certain teleoptiles appear downy and are referred to as semi- plumes, but true down feathers, or plumulae, are absent. Also, the neossoptiles which make up the downy coat of the newly hatched chick are usually attached to the ends of the following juvenile plumage, or mesoptiles. The teleoptiles are the typical feathers of the adult. An outstanding characteristic of the feathers of the grouse is the aftershaft — a second shaft bearing most of the typical feather parts and attached near the base of the main shaft. It is best observed among the contour feathers. * By John E. Trainer. A See Appendix, p. 741. r\ LIBRARY ]^\ ^V MASS. X«S/ ^< -V J ^-E 'A ■■■{■■ X( ^ ' '«r tr ■'• 60 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE The feathers of a female grouse (teleoptiles and seniiplunies) were counted as they were plucked and found to number 4,342*. In this species there are ten primaries and about 15 secondaries. The tail is normally composed of 18 but from 16 to 20 have been recorded. Each year grouse grow a series of rod-like appendages as a fringe along the sides of the toes in the fall and shed them the following spring. The bird is thus provided with "snowshoes" for the winter. These rods are derived from the skin and are thus closely allied to the feathers. PHYSIOLOGY'^ When winter comes man puts on heavier clothing and lights a fire in his furnace. Many northern mammals grow a thicker coat of hair or go into hibernation. Grouse seek shelter in thick evergreen clumps and fluff out their feathers to provide insulating air spaces. Main- tenance of body temperature is of paramount importance to the well-being of all warm-blooded animals. Yet means of regulating heat loss represent oiilv mif side of the picture. The manufacture of heat through the burning of food materiuls ii' the ImkU must also be carried on. Unless the two are suitably coordinated an animal cannot survive long. One should realize too that wild species, unlike civilized man. do not have access to arti- ficial heat nor can they put on extra clothing at will. This is particularly true of birds which do not even grow a heavier coat of feathers. They must, therefore, rely to a much greater degree on producing within their own Iiodios the added warmth necessary in cold weather. Production of body heat is dependent upon the proper functioning of the various internal organs — in other words, the animal's physiology — as well as upon adequate shelter and food. These vital processes are !iot static. Rather they undergo constant adjustment to changes in the activity of the animal and the environment in which it lives, as well as in the amount and quality of food eaten, the presence of disease and other influences. * Lone. W. H. (unpublinhrd) foiinj tlie frathcri n( 24 adul! ermine In nvrraRp 7 per cent nl llii- total iiiiti.il wfichl. A fly William !I. Long, PHYSIOLOGY 61 There are, however, limits to the adjustment which an animal can make. When environ- mental conditions become more severe than it can cope with, death results. But even though survival is not endangered, changes in the conditions to which an animal is subjected affect its vigor and behavior. The varying cumulative effect of such forces at different places undoubtedly is a major factor in determining the extent of the distribution of a species. Similarly, at different points within its range these relationships govern to a consid- erable degree seasonal need for various types of food and shelter. Again, severe conditions during one year may so lower an animal's vitality that it is unable to recuperate in time to survive the next. Therefore, a knowledge of the principal environmental influences involved and of how they affect the species in which one is interested will aid both layman and technician in recog- nizing habitat deficiencies. Likewise the game manager will be better able to devise effec- tive development measures. Only recently, however, have detailed studies of these relationships been undertaken deal- ing with game species. The data presented herewith are the first pertaining to the ruffed grouse*. With respect to this species the basic environmental forces affecting it physiologically are air temperature, air movement, radiation of heat, evaporation, humidity and light. Factors such as shelter and food merely modify a bird's response to the other niore fundamental influences. A GROUSE WITH ITS FEATHERS FLUFFED OUT TO PROVIDE INSULATION AGAINST THE COLD In measuring the effect on the bird of such conditions or combinations, the jirimary indices, in addition to survival itself, are body temperature, respiration rate and heart rate. But before significant variations can be recognized one must know what the normal is. Initial experiments were jilanned accordingly. Then, following these, others were conducted to determine the effect of various environmental changes. In connection with the data secured, however, one must bear in mind that the birds used were from hand-raised stock and may not be entirely representative of wild grouse. They were also all adult. * A more detailed accuiinl of these experiments is included in the Appendix, p. 749. 62 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE It was found that the average body temperature of the birds used was just over 107°F. and that it varied slightly with the air temperature and with activity. The data also demonstrated a regular fluctuation each 24 hours, in which the temperature of the birds was highest between 2:30 p. m. and 5:30 p. m. and lowest between 11:30 p. m. and 2:30 a. m. Wetting the birds with a water spray in simulation of rainfall caused their body temperature to decrease sharply. The colder the water as well as the surrounding air, the greater the degree of drop and the slower the rate of recovery. The effect was also much more severe on three speci- mens afflicted with an intestinal disorder. The average rate of breathing among inactive grouse was nearly 63 respirations per minute. This rate increased with the activity of the bird, as well as with the body tempera- ture. It appears, however, to be a less significant indication of relative vitality than either body temperature or heart rate. The average heart rate of inactive grouse was found to be about 342 beats per minute. Like respiration rate it varied with the bird's temperature, but there was a tendency for it to become more rapid as the en\ir<)nnient became colder. When the birds became excited a pronounced and almost instantaneous increase occurred. To study the effect of starvation on the bird, 12 grouse were held for nine days with- out food. While this did not markedly affect the general level of body temperature until the last day, it did cause a drop in the low point of the 24-hour cycle each night. Since these birds were held at a constant air temperature this drop would be greater under natural condi- tions where the air becomes colder at this time of night. Thus birds without adequate food would have greater difficulty than well-fed ones in preventing excessive reduction of body heat. Furthermore, the fact that these birds did maintain their body temperature, especialh when living at low air temperatures, demonstrates that such conditions force birds to draw on their reserve in order to compensate. In addition to the above, six grouse were allowed neither food nor water. After the 48th hour under these conditions their temperature-regulating mechanism broke down and in sev- eral cases death resulted. A definite daily need for water is thus apparent. WINTER CROU.se DROPPINGS (LEFT) RESULTING KKOM GOAKSE, IlK.llLV FIBROUS FOODS AND SPRING DROPPINGS (right) PRODUCED AFTER THE BIRDS BEGIN TO FEED ON FRESH GREEN MATERIAL PSYCHOLOGY AND BEHAVIOR 63 Under starvation conditions birds which were heavier at the start maintained their weight better than did lighter individuals. Weight loss was greater for all birds at night than during the day time. Under normal conditions food was found to pass more slowly through the alimentary tract as the environmental temperature dropped. Finally an effort was made to determine the amount of food needed by an adult grouse to maintain its weight under average living conditions. Since differences in composition between various foods cause their fuel values to differ, a standard unit of measurement, the calorie, was used for comparison. It was found that, for the birds studied in this experiment, a food consumption having a value of 78 calories per day should result in their neither gaining nor losing weight at an environmental temperature of 65 -F. when not subjected to exertion. PSYCHOLOGY AND BEHAVIOR* Each activity of a living organism is a component of its behavior. Reactions are largely the result of an animal's physiological and psycholopicai makeup in response to external stimulation coming from the habitat in which it lives. Therefore, a wildlife manager must understand and take into consideration as many of the patterns of behavior as he can if he is to take full advantage of the possibilities of increasing wildlife abundance through devel- oping and managing its environment. Yet, with most game species, grouse included, very little is known about these basic relationships. The Investigation has had an abundant opportunity to note grouse behavior in the wild and in captivity. Consideration of the pertinent observations gathered therefrom falls logi- cally into two categories. Either one may desci ibe the actions of the birds under various circumstances primarily from the standpoint of a disinterested spectator, as in Chapter V, or one may go beyond that and attempt to understand such behavior in terms of its significance in the life of the bird. The psychological aspect of the latter approach seems sufficiently distinct to warrant special alli-nlidn lure. Two such behavior patterns have become clearly enough defined to merit description. The first is the social order; the second, that of mating behavior. It should be recognized at the outset that it is difficult and ofttimes misleading to attempt an analysis of animal psychology, based on behavior, because of the inevitable tendency to interpret reactions in terms of their human counterparts. Likewise, pitfalls beset one in trying to evaluate observations made of captive individuals, which of necessity have been the basis of much of this discussion with respect to grouse. Realizing this, no suggestion as to their degree of applicability to birds in the wild is contemplated here for. in doing so. one may be guilty of ascribing to such birds responses possibly induced in their hand-raised cousins to some extent by the artificial conditions under which they are held. On the other hand, the reactions here described are basic and almost certainly are to be found among wild birds as well as among those held in captivity. Let us then depict them here. The Social Order The ruffed grouse may not normally be considered a gregarious or social bird. Yet from the time the chicks are first hatched until well into the fall, numbers of individuals are found together either in the loose association of the brood or, very occasionally, in so-called * By Gardiner Bump. f)l THE SPEC.IES~ITS TAXOSOVY. n.4\GE. RIOLOGY. & ECONOMIC IMPORTANCE "packs", the latter probably representing several broods or groups of iiuiividuals gathered together at a particularly favorable feeding site. Likewise, throughout the late fall and winter more than one bird may resort to as small an area as a cluni]) of conifers for night roosting, thus bringing it into relatively close contact witli others of its kind. But it is to captivity, where the birds are grouped in pens, that one must turn for examples of the social order which, were we more skilled in making observations, might be recognized as also existing whenever grouse congregate. At the end of each brooding season at the Research Center, the birds of the year, along with the breeders, are placed in wintering pens where they remain as groups until the begin- ning of the following breeding season. Here, as has been described with poultry" there is soon established a rigid and demanding social hierarchy. In ruffed grouse, where the sexes are relatively similar in appearance, the basis of this social hierarchy apparently lies, not primarily in the age, sex or weight of the indi\idual birds, but rather in the vigor of the in- dividual and the degree to which it is able to dominate its penmates. Success in fighting, in bluff and, to a lesser extent, in display, such as strutting, rapidly establishes a definite social order in which each, save for the bottom bird, exercises a dominance over its less fortunate penmates. Except for the ranking bird, each is in turn dominated by its physiologically and psychologically stronger companions. THROUGH HOHlliNU AiNU BLLKHiNG A DEFIMTE SOCIAL OKDKK IS KSTAlil.lMI KD AMONG GROUSE, BOTH IN THE WILD AND IN CAPTIVITY In captivity, serious consequences may often follow this process of establishing such social rank. A bird, defeated in one or another of the frequent combats that take place, is occa- sionally pursued by the victor into a corner of the pen and there pecked on the head until the skull, in extreme cases, is laid bare. Once completely subjugated and without the bene- fit of self-confidence, if it does not die of mechanical injury or stanation. it soon becomes PSYCHOLOGY AND BEHAVIOR 65 the object of attack by most other birds in the enclosure. If, on the other hand, the combat ends in the weaker bird edging away without being pursued, meanwhile recognizing the victor as the dominant individual, it may in turn be successful in combats with others, thus estab- lishing its own position without serious harm to itself. Although the order is not directly determined by sex or age, the fact that the heavier and larger birds are principally males over one year old usually results in a larger proportion of these birds occupying high ranking positions of dominance when groups comprising all ages and sexes are penned together. Since those low in the scale must keep out of the way of their stronger companions, the practice of segregating males from females and, occasionally, adults from birds of the year, has grown up. Even in such groups, however, a dominance order soon develops. It is interesting to note that, should a dominant bird in one pen be placed in another, it must establish its rank among its new companions by the same method as heretofore described. In this case, however, the new arrival is at an added disadvantage in that the resident birds have also established a feeling of ownership of the territory represented by the pen. Apparently the newcomer recognizes this and is, therefore, placed at a psychological disadvantage at the start. It is for much the same reason that a bird once removed from an enclosure and later returned thereto, does not always reestablish its old position in the social scale. Also, if a vigorous bird, high in the social order, is weakened by disease or some other factor, it will be forced by its companions to assume a lower position in the hierarchy. It is such considerations as these that have given rise to the practice of shifting a bird in captivity from one pen to another when it becomes so strongly dominant as to upset the social balance of the pen. Translated in terms of relationships among wild grouse, it is possible that the existence of a strong dominance complex, particularly among the males, represents one of the important factors limiting the numbers which will occupy a given habitat. One likewise finds here a possible explanation for the breakup of a brood in the fall for, at least in captivity, the young birds, as early as the first half of September, begin the conflicts that lead to the establishment of a social order. Mating Behavior Aside from dominance, the only other behavior pattern which is sufiBciently distinctive to warrant special consideration outside of the chapter on General Habits, occurs during the mating period in response to the marked physiological and psychological changes associated with the reproductive cycle. Here, too, one must turn largely to observations on captive grouse for a knowledge of reactions and an interpretation of their meaning. Watching grouse activities throughout the spring and early summer, one is impressed by the number and variety of reactions to be obser\'ed. With close study, however, they will be found to fall logically into three phases as here described. Strutting Phase In the pens at the Research Center, where males and females are by necessity placed together most of the year, the males show signs of aggressiveness towards their penmates of either sex as early as the first of March. Though they may have been strutting* on warm days throughout the winter, they now display more frequently and with greater vigor, even in the * See Chapter V, p. 282. 66 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE absence of the female. Apparently they feel the need of making themselves appear conspic- uous and formidable, possibly as an aid in maintaining their dominance complex. In fact, Alien'" has termed this the "intimidation dispia\"'. As the season progresses and excitation mounts, a locomotive-like hissing and violent head .shaking are added to the strutting performance with increasing frequency. Occasionally, they may select a high point and drum. These, it would seem, are the first steps in their spring breeding pattern. Gardiner Hump THE STRUTTING MAI.K SEEMS BOTH TO ATTR.ACT AND TO INTIMIDATE THE FEMALE At the height of this first phase, so strong is the physiological stimulus that, in captivity, full display may often be inspired by the presence of man or of many other objects not familiarly associated with the enclosure. In fact, almost any change in the innnediate environ- ment at this time will result in a display. In securing recordings of grouse calls, for instance, the male would frequently respond in this manner to the placement of the microphone in the pen. Throughout this phase, the male seems preoccupied with establishing and maintaining his territory, as well as with display. At first he accepts the enforced presence of the female; later he is increasingly stimulated by her. She, in turn, evidences little apparent interest in his reactions except for a desire to keep out of his way. If display becomes too vigorous, she may spend considerable time hiding or on a perch in an attempt to escape his over-zealous attentions. Tliough in the latter part of this phase, the male may be physiologically ready to mate, the majority of the females have not as yet reached this stage. PSYCHOLOGY AND BEHAVIOR 67 Gentle Phase It is the second, or gentle, phase of the mating cycle that, in the male, is most distinct and easiest to recognize. At its height he seems to abandon for a few days all interest in terri- tory and in striking display. He is subdued and relatively inactive, preferring to sit still or to DURING THE '"GENTLE PHASE Gardiner Bump BOTH SE\ES AKE QIIET AND M HDl KU IN ACTION follow the female at a slow, careful walk, meanwhile twitching his head and neck slightly. Occasionally he repeats a call somewhat similar to that of a female singing on the nest. His whole attitude is the epitome of gentleness. Often he follows close after the female and, when she pauses, lightly places one foot on her tail or back, meanwhile pecking her softly. When she moves away he follows slowly, almost as though in a trance. For her part, the female perhaps recognizes that her male, abandoning his pugnaciousness for the moment, is in a gentle mood. H she is in the period of sexual desire or oestrus, she may squat and assume the mating posture*, whereupon mating may take place. But, because this second phase seldom lasts, in captivity at least, more than a week, coition has been more frequently observed at the Center during the last part of the preceding phase, and the first part of the phase to follow. More often, she is inclined to lead him here and there about the pen, ofttimes singing and occupying her nest box, interested in his attentions but not yet ready to yield. It is almost as though, for a brief time, she recognized some bond of mutual under- standing. Fighting Phase Transition to the third or fighting phase may come swiftly. In it, for a time, the male may return to the strutting display characteristic of the first period, perhaps by way of finding an outlet for the intense energy and stimulation so evident in him at this time. Gradually, how- ever, as interest in the female wanes, he develops a strong pugnaciousness in defense of his * See Chapler V. p. 267. (,[', THE SPECIES— ITS TAXONOMY. RANGE. BIOLOGY. & ECONOMIC IMPORTANCE pen against all comers save only those of whom he is mortally afraid. Assuming a charac- teristic fighting attitude with head low, he will follow the attendant, as the latter moves around the pen. and try determinedly to get out and at him. Should the pen he entered, the bird will fight with surprising vigor and viciousness. The same holds if another male is intro- duced. Even when a strutting hird is approached at this period, the change to fighting pose is commonly accomplished in from five to 15 seconds. Following the disappearance of the intruder, the bird may resume his strutting or carry on various pen activities, meanwhile keep- ing a sharp lookout for the next visitor. As spring merges into summer the male gradually abandons strutting altogether and not long thereafter may become incapable of being stimu- lated to the point where mating can take place. The fact that he exhibits the fighting reaction rather than strutting is, however, by no means a good indication that he is not physiologi- cally able to mate. k^^. '■■trdiner Bump Willi. K IN THK FIGHTING PHASK TIIK MALE EXHIBITS CONSIDERABLE PUGNACIOUSNESS IN DEFENSE OF HIS TFIUUTORY For her part, the female, during her third phase, exhibits little interest in the male except to keep out of his way. This is her period of egg production and considerable lime is accord- ingly spent on and about the nest. Allen'" has given an excellent description of her behavior at this time. Beyond the third phase the reactions of the male are characterized by a complete lack of interest in his mate except as an individual to be dominated if occasion demands. For her part, she reciprocates in kind by going about the business of incubating her eggs and raising her brood. Her actions throughout this period are described in the chapter on General Habits. In interpreting certain of the reactions on which recognition of the three phases here described is based, one experiences real difficulty. In studying breeding behavior, Allen con- cluded that grouse did not recognize sex and described in the male a recurring mating rhythm involving physiological and psychological readiness to male. The present authors, in the PSYCHOLOGY AND BEHAVIOR 69 course of experiments, have many times observed the reactions on which Allen's conclusions were based. Complete records taken twice a day of the occurrence of each phase in all the breeding grouse at the Research Center over a period of two spring seasons have also been kept (table 7). Analyzing these, a different interpretation is here suggested. Let us consider for a moment the matter of sex recognition. Unquestionably male grouse may display to a bird of either sex or even to unfamiliar objects, as previously mentioned, during the height of their display phase. At other times the presence of a female seems more likely to stimulate this response by the male. Likewise, place both a female and a male with a displaying cock for a few minutes and, providing one bird is not too over-dominant, the latter will often intensify its strutting in the direction of the female and exhibit a desire to fight the male. Two cocks do not commonly display to each other. Nevertheless, as suggested, one function of strutting may be intimidation. No evidence has been noted of a recurrent sex rhythm in the male. True, as described by Allen, there are times during all of the three phases when most male grouse will mate with another bird if it be posturing and other days when such birds will not mate at all. In the first place, strutting, to which, at the height of excitation, the hissing and headshaking reactions are added, is ])redominant. The second phase, characterized by gentleness and head-twitching, follows in turn, to be superseded in a short time by the strutting and chasing characteristic TABLE 7. A GENRRAIJZED BREEDING BEH W lOR PXTTERN COVERING ADULT MALE RUFFED GROUSE IN CAPTIVITY COMPILED FROM A STUDY OF 78 INDIVIDUXL PATTERNS Hour April 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 SAM s s s N S S S S H S S S S 11 H S S s 11 II n s H H S H H H S C S 11 M S S C S S C S H 5 p M .S 8 A.M. 5 P.M. May _L 10 11 IS 16 17 18 19 20 21 22 23 23 StruttiiiK Hfiad-twilchinjr C = Chasing Activity not cliuracleristic of breodinp pi'rioil of the third phase. If one charts for all the males the occurrence of each phase based on characteristic behavior, there appears to be little tendency to pass through these more than once in a single season. A bird, of course, does not change from one phase to another overnight. During the interval, which may last for a week or more, one finds the behavior characteristic of both phases exhibited many times. Once the bird is well into a phase, however, except for strut- ting, one does not commonly observe the behavior characteristic of the next stage until it is approached. Perhaps the situation, as it seems to the authors, can be clarified by considering that male grouse in ca])ti\itv normally are in a state of physiological readiness to mate during the latter part of the first j)hase, throughout the second and the first part of the third phase. The act is not normally carried to completion, however, unless the male is externally stimulated above a point one might call the threshold of mating. 70 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE Light, heat, air movement, food, mood of the female, are some of the factors varying! from hour to hour which may serve to stimulate a grouse psychologically to a point above this threshold. Properly stimulated, he may attempt coition in the morning: change the external conditions and he mav avoid the act in the afternoon. L«:k<«2«tiL«^'»iA& Gardinrr Bump ROMEO AND JULIET EGGS 71 To prove that he is physiologically ready, however, in several such cases the birds, under manual stimulations, have ejected sperm. Females not previously bred, artificially insem- inated with this, have in turn produced fertile eggs. Also, given the right conditions, males have been successfully mated with three difFerent females in the course of one day. Perhaps then changes in external stimuli, rather than a recurring sex rhythm, cause a male grouse to be ready to mate at irregular intervals throughout the breeding period and to refuse coition in the meantime. EGGS The ruffed grouse most frequently lays from nine to 14 eggs, although both larger and smaller numbers are not uncommon*, the latter usually indicating a renesting bird. The eggs are ovate in form. In color, they vary from milky white to cinnamon buff, usually plain but often spotted with reddish or drab. The shell is smooth with a slight gloss. As incubation progresses, they often become considerably stained with dirty, brownish blotches. A (,H()LF or TYPICAL GKOLSE EGGS * See Chapler VIU, p. 360. 72 THE SPECIES— ITS TAXONOMY, RANGE. BIOLOGY. & ECONOMIC IMP0RT.4NCE During the seasons of 1937 and 1938, measurements were recorded for 366 grouse eggs in 30 nests on the Connecticut Hill area. The average of this group was 39 x 29 millimeters. The largest egg measured 49 x 33, although the next largest were 42 x 31 and 39 x 32 milli- meters respectively. The smallest was 36 x 28 millimeters. Regarding size, Bendire" states: "The average measurement of forty-four specimens (of B. u. umbellus) in the United States National Museum collection is 38.5 by 30 millimeters, the largest egg of the series measuring 40 by 32. the smallest 33 by 25 millimeters." The type clutch of this group was secured near Harper's Ferry, West Virginia, but no locality for the others is given. Bent" records the average of seventy-three eggs as 38.9 by 29.6 millimeters. This group seems to have included that reported by Bendire since the extremes are identical except for maximum length which in the latter was 42.7 millimeters. In Maine, a clutch of twelve eggs measured by Knight^" averaged 37.8 x 29.4 milli- meters*, while a series of nine from Maryland"" (apparently not from the same clutch) averaged 41.4 x 29.7 millimeters*. Occasionally abnormally small eggs are encountered and the Investigation has recorded one double yolked egg. According to Bent" the eggs of the various subspecies are indistinguishable, akhough those of sabirti show a possible tendency to be "slightly darker in color and somewhat more often spotted." EMBRYOLOGY^ The question of which came first, the grouse or the egg. may not be pertinent, but to what degree the egg controls the future destiny of the grouse is certainly of utmost importance. An investigation of the embryonic stage, therefore, was one of the first natural steps in trying to discover the underlying factors which cause the fluctuations in abundance of this unpre- dictable bird. PREVIf;\V OF THK PROBLEMS With grouse, however, the saying "Don't count your chickens before they hatch" should include "and live". Both the hatchability of the eggs and the livability of the chicks are vital to maintaining population levels. Furthermore, they are closely related. In studying the eggs with respect to their bearing on these factors, many other problems had to be taken into consideration also and attempts made to solve them, either in whole or in part. Among these was the quality of the eggs and whether or not it varied between years or even during the same season. Also involved were the influence of climatic or solar changes, the function of essential food ingredients and variations in their supply and the effects of dif- ferences in egg conformation on both the growth of the embryo and the mortality of the chicks. HalchabilUy Will the egg hatch? If it has been fertilized, it may. But there is no way to find out • Thrm. mcaMUrrmrlilH were Kivrn by the autllnrs in incheii but liavt- liiTii riiiitrrteil In iiiilliiiicteiK fttr citiiiparAbilily. A The rxperiini-nli* dinrtiHsfil under this headinE were carried on by Dr. A. I.. Komannff of Cornell Vniversily, collaborating willi the InTeatigalion. EMBRYOLOGY 73 except by incubating it and awaiting developments in the embryo. Among eggs from wild grouse the proportion hatching successfully has been consistently high and variations apparently have had little relationship to fluctuations in the number of birds. Among those from captive grouse the proportion has been somewhat less but the difference has been slight. Livability Will the chick live? This is a more difficult problem. The high mortality rate of the young birds even after a successful hatch has been a primary factor in regulating the size of fall populations. Yet what causes it cannot be determined from field data alone. Both the quality of the egg and the environment into which the chick is born, play major roles. Labora- tory and field research must be coordinated in order to find out what those roles are. Realizing this, the Investigation undertook a five-year study, from 1936 through 1940, to determine, as far as possible, how much the inherent qualities of the egg influenced the growth of the embryro and the viability of the chick. In all. 398 wild ruffed grouse eggs from 44 clutches, collected near Ithaca, N. Y., and 189 newly hatched chicks were analyz- ed. The study included the general development and structure of the eggs, their physico-chem- ical properties, annual changes in quality, develojiment of the embryo and embryonic mem- branes, and embryonic mortality. It also included the chemical composition of hatched chicks and its annual changes. To date, the Investigation has not carried on extensive enough studies of the rest of the prob- lems to report findings. The influence, if any. of climatic or solar changes on concentration of essential food ingredients, as minerals and vitamins in plants and even insects, the effects of food supply on egg development, how weather reacts on the quality of the eggs, and the pos- sibility that variations in egg conformation may retard growth of the embryo and help deter- mine livability, are still problems for the future. Development of Grouse Eggs Grouse eggs in their development resemble in many respects those of other gallinaceous birds, such as pheasant and quail. The incubation period is normally about 23.5 days. It is only a few hours shorter than that ol ])heasaiits and about one day longer than that of quail"". From the weight of eggs and birds at hatching, it is also evident that the growth rate of the grouse embryo is somewhere between the growth rates of pheasant and quail embryos'"". Structure of the Egg In structure the fresh grouse egg is, in general, similar to the eggs of other birds. It consists of yolk, albumen, shell and blastoderm ( figure 41 . The stratification of yellow- and white yolk corresponds to the day and night growth of the yolk in the ovary. The four layers of albumen, with two cord-like attachments at the opposite poles of the yolk, oc- cupy nearly two-thirds of the entire egg. except for a small space at the large end known as the air-sac. Tlie inner surface of the egg is lined with two parchment-like membranes. The blastoderm, appearing like a small whitish speck on the upper surface of the yolk, is the seat of the future embryo. Physico-chemical Properties of the Egg Table 8 gives the average total weight, shape, breaking strength, weight and thickness of 74 THE SPECIES ITS TAXONOMY, RANGE. BIOLOGY, & ECONOMIC IMPORTANCE eggshell, and the thickness of shell membrane. All these, including the total ash and calcium content of the eggshell, facilitate the functioning of the specific characteristic properties of the egg of this species and are called its values. They are noticeably different in many respects from those of the eggs of pheasant, quail and other domesticated birds'^. BLASTODERM ALLANTOIS ALBUMEN YOLK ALBUMEN YOLK SAC FRESH EGG ELEVEN DAY EMBRYO FIGURE 4. STRUCTURE OF THE FRESH AND OF THE DEVELOPING GROUSE EGG On the other hand, some of these values, when corrected for the size of the egg, show striking similarities. The chemical composition of eggshell and possibly of egg contents, on a percentage basis, are much the same as in the eggs of other species, especially of pheasant and quail. The variation in relative values among individual eggs of the same species is fre- quently greater than the variation between the averages for different species. There was little difference in physical properties and chemical composition between grouse eggs from birds in the wild and from those in confinement. The eggs of wild birds had somewhat stronger and tiiicker shells, with greater content of total ash and calcium. It would seem from this that artificial environment and possibly the lack of natural foods have modified the shell slrintiirc. There was a tendency for the eggs of captive birds to become pro- gressively smaller, more elongated, and to have lliiriiirr. liiiiilcr. and more easily broken shells, the later in the season ihev were laid. TABLE 8. PHYSICO-CIIHMICAL PKUl'KUTIKS OK CHOUSE EGGS COLLECTED KIlOiM THE ITHACA, NEW YORK, REGION 19.S6-1910 *ERg8 incubated a few days bcfora the time of delivery to the laboratory weighed somewhat leaa than they should at the oorreapondinff period in the wild. EMBRYOLOGY 75 Annual Changes in Egg Quality So far, only the changes in various physical and chemical properties of the eggshell have been determined. Data for the composition of the egg contents are not sufficient to warrant reporting. It was established that there are differences in shell quality, year by year and. during any one season, between clutches and even between eggs in a dutch. For example, figure 5 shows annual variations in eggshell thickness for the five-year period of the study, the year- ly averages for each clutch and the different thickness of each eggshell. The average thick- ness during 1936 and 1940 was higher than fur the intervening years. Yet each year had some clutches with high and others with low averages, while individual eggs varied greatly. _l UJ I (/) O O UJ u O o o tn CO UJ z o I I- 023 021 019 017 015 013 01 I? 1936 I I I I 2 3 •5? 1937 12 3 4 5 . . • INDIVIDUAL VALUES O AVERAGE PER CLUTCH • I I • i . 1938 I I I I I I2345G7 123 NDIVIDUAL CLUTCHES I • o » : o i . I 1940 I I I I I I I I I I I I 234 56789 10 II FIGURE 5. ANNUAL CHANGES IN SHELL THICKNESS OF GROUSE EGGS COLLECTED FROM THE ITHACA, NEW YORK. REGION — 1936-1940 Since hatchabilily is consistently high, a change in eggshell thirkncss seems of little sig- nificance. However, future study of thickness and composition of the shells of eggs which had hatched, as compared with those in the clutch which did not. may give some new evidence as to how these qualities influence hatching. Their effect on livability may also be determined by checking tlicm against the sulisequent mortality of the chicks. More time and greater numlters than have been availalile fur ibis study are required to make tlic data r-i')io ObservntionB Avnragft value in prams T.ivp wpipjht 1.3.0 \\t'i"lit witlioul volk 11.1 \\i'i;.'lit (if yiilk siic with contents. l)rv wt'ifjht 1..-.: 0 92 0.21 Calcium (CaO) content 0.078 to the body weight, are comparable to those of other game birds, especially jiheasants and quail. On the other hand, the pro])ortion of unabsorbed yolk to the body weight at hatch- GROWTH AND DEVELOPMENT 77 ing. while quite similar for the three species of game is, however, much smaller than in some domesticated hirds. such as poultry. This means that grouse, after hatching, can survive only a relatively short time without food. The chemical analysis of newly hatched grouse showed some annual changes during the five-year period. However, the occurrence of a few highly variable values frequently inter- fered with the analysis of these chemicals. For example, in most of the clutches the weight of a chick without the yolk was the lowest in the year 1937 (figure 6). but the average to- tal weight was nearly the same as in the other years. 14 ? o z 13 - 1- ■D 1 1 o T H $ 10 u (/) O 9 a. o u. fl o 1- 7 1936 J L I 1937 12 3 4 J L J_ _L 4: • 6 I t • • • INDIVIDUAL WEIGHTS o AVERAGE PER CLUTCH 1938 j_ _L _L o I 1940 J L I 2 3 1234 1234567 123 INDIVIDUAL CLUTCHES FIGURE 6. ANNUAL CHANGES IN THE WEIGHT. WITHOUT YOLK SAC, OF NEWLY HATCHED GROUSE COLLECTED FROM THE ITHACA, NEW YORK, REGION — 1936-1910 Besides these variations, it is also possible that the selected measurements were not signifi- cant. More delicate measurements of physical and chemical phenomena of embryonic devel- opment may. however, disclose greater annual differences in embryonic metabolic activity. It is a question whether deviation in such activity could have been responsible for the lower viabilit> atui early death of certain embryos, and thereby be one of the causes of low grouse abundance. GROWTH AND DEVELOPMENT* The phenomenon of changing the protective coat coincident with growth of animals is ac- complished in many interesting ways. Fishes, for instance, retain the covering in which they are born but the scales increase in size and afford complete protection throughout life as » By Earl R. Holm. 7y THE SPECIES ns TA\0\UMY, RAXGE, BIOLOGY, & ECONOMIC IMPORTANCE may be noted from their annual growth rings. In certain reptiles the comijlcte skin is shed at intervals during the period of growth. Among birds, a series of regular feather moults main- tains the necessary protection to the body not only by compensating for growth but also by adequately covering the surface to a greater or lesser degree in accordance with seasonal changes in temperature. Apparently the primar) function of feathering is protection. This coat is not only a tem- perature regulating device offsetting the effects of the elements, but also affords protection fiiini natural enemies. Probably everyone who has hunted grouse has had the experience of picking up a bird, only to have it make its escape leaving but a handful of feathers. The shedding of feathers in this way seems to be controlled by voluntary muscular action and is undoubtfdlv utilized in escaping from predators. The replacement of feathers lost in this way or by other accidents probably explains many of the variations in the normal pattern of tlie moult. Much has already been written on the moult and sequence of plumage among birds. Dwight"" was one of the first to disoel many of the old theories which persisted even into the present century. His studies of the passerines as well as certain of the gallinaceous species represented a major advance in our knowledge of this subject. Considerable work, too, has been done on domestic fowl. Warren and Gordon""' carefully noted the number of days required for the replacement of the flight feathers in certain breeds of poultry. Other studies were conducted l)y Radi and Warren'" and by Marble'"' "'■ specifically to determine the relation- ship between feathering and growth as well as egg production. More recently Host"", work- ing willi a close relative of our native grouse, found light to be the main factor controll- ing the development of spring and summer plumages in ptarmigan. Many other researchers have done equally important work on other phases of the subject. In view of the forego- ing, the emphasis in this discussion is placed on the sequence of the moult in the ruffed grouse as an aid in age identification*. Students as well as research agencies working with immature gallinaceous game birds have often been at a loss to estimate even approximately the age of specimens with which they were dealing. Sportsmen, too. would like to be able to differentiate between birds of the year and mature specimens a full year old or older. Such knowledge ofttimes might be quite convenient for when the game is divided at the end of the day one could gener- ously give the older heavier birds to his companinn and retain the younger more tender ones for himself. Eaih would be well satisfied since both woidd feel sure that they best- ed in the deal. By studying the feather patterns and sequence of llie moult, paitirularly in the flight feathers, it appears that age can be determined within relatively narrow limits when dealing with average birds. Since the feather patterns are discussed in detail in the Appendix under the Pterylography of the Ruffed (Jrouse. il will suffice to say here that the arrangement of the feathers as well as their rcplai I'nicnt during llic mcmlt follows an orderly and regular sequence. This is con- trary to the once pojjular cdtK cpliipn that the feathers are shed like the leaves from a tree and re])laced all at once like buds bursting in the spring. Sequence of Moii.t ano Feather Succession The sequence of the moidt may be di\ idod into three general plumage classifications, na- tal, juvenile and adult. It should be borne in mind, however, that there is no definite line of * Till- fiillowitiK (lifcussiim in baiir.) itii llir tuidy Irnm tinlrh:iit: h> muliirir\ of a Ijrg.- (..-rirs of gr«ii»r rrurni at tlic Rrscarcll Crnlcr, Delmar, N. Y. GROWTH AND DEVELOPMENT 79 demarkation between one and the other since the moult is in constant progress until the bird reaches maturity. Even while the chick is in its natal down* the flight feathers or remiges are developing. In like manner when the juvenile plumage covers most of the body, the na- tal down is still present in certain feather tracts while the mature plumage is developing in others. In certain instances the natal down is replaced directly by feathers identical in pat- tern and structure to those of the mature plumage. Thus the intermediate juvenile feather does not occur in this sequence. Natal Plumage At the time of hatching, the body of the grouse chick is almost completely covered with feathers that are popularly called down.* All of the spaces, i.e., areas between the regular feather tracts, are bare although they are obscured by the fluffy down. The most conspic- Doug Finch THREE GROUSE CHICKS SIX HOURS AFTER HATCHING uous of these are the areas over the lungs (which are directly beneath the folded wings), the web of the wings and also the sides of the abdomen. The down as well as some of the flight feathers begins its development during the early stages of incubation of the egg and growth apparently is uninterrupted by hatching. * Tcrhiiiinlly the featluTs ii( tliis* pliimuge are neossMptiles. true il.iHti fi-alhers ( iiluimilae I being ahs.'nt. p. 59. 80 THE SFECIES~ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE After the chick has dried off and takes its first furtive peek at the world from under the wing of the mother, seven primaries may be counted. The longest of these usually has approximately one-half inch exposed and a few will have about one-eighth inch of the feather tip protruding from the sheath. None of the other three which make up the complement are visible. While the average number of secondaries is 15, only nine are visible in the day- old chick. Most of these have the downy tuft appended. The tail is represented by fuzz in the form of longer drooping feathers. Juvenile Plumage The juvenile feathers begin to appear immediately after hatching led by two or three scap- ulars which may be noted at four days of age. The flight feathers which are present at hatch- ing, however, are functionally juvenile. When it occurs, the intermediate plumage represents a transition stage wherein the size and number of the feathers are adapted to and conform with the progressive increase in size of the bird. The feather which replaces the down develops in the same follicle and as it emerges the latter is forced ahead of it. The downy tuft remains appended to the tip of the sheath of the new feather. Unless it is broken off it remains attached until the sheath opens ex- posing the lip of the developing feather. Thus the down with its elongated shaft and the new juvenile supplement each other to afford the necessary protective covering for the body. It is evident, from studying feather succession, that the replacement follows a definite pattern designed to afford the chick the greatest protection compatible with its habits at a given age. Hence, the lungs, which are probably the most vulnerable organs of the young chick, are pro- tected by the flight feathers, wings and rapid development of the scapulars (figure 7l. 1 he latter merge with the back feathers and close over the back of the bird at about three weeks of age. This coincides with the time it is being weaned from brooding by the mother. Coin- cident with this growth, the feathers on the sides appear although these are obscured in the early stages by the dense down. It is apparent that when the chick snuggles into the breast feath- ers of the mother for warmth, the naked area over the lungs may be exposed to the maxi- mum heat from her body. \\ hile there are certain exceptions, the pattern of succession in general is such that the first feathers which appear cover and protect the base of those subsequently de\eloping. In line with this, the emergence of the scapulars progresses tnuaid the rear from a point op- posite the bend of the folded wing. These feathers also dext-lop laterally to merge with those of the back at the base of the nape. Those of the sides develop from front to rear and downward to cover the bases of the flank and upper IkIK feathers respectixely. The feath- ers in the uj)p<'r rump region appear first, followed li\ those toward the rear which cover the tail coverts. .Similarly the tail coverts attain sufficient length to cover the base of the tail before the pin feathers in the latter appear. The flank feathers come in before those of the thigh, while the appearance of the latter commences at the up|)er end of the leg. progress- ing downward along the tibia and tarsus. These are feathered on the rear and front and finally become closed in at the sides. On the belly, the succession has a tendency to spread ill all directions. A bird at three weeks of age has a triangle of down extending backward from the forward point of the keel, with its apex about midway to the pel\ is. This triangle i-i bounded laterally by a fringe of juveniles. At the forward part of the belly this fringe consists of onlv one feather on each side. Following to the rear on each side a pair appear, then three rows and finally four rows where the two sides join at the midpoint on the keel. Eight rows of feathers extend from here to the rear point of the keel. In the belly area the GROWTH AND DEVELOPMENT 81 feathering progresses up toward the sides and flanks and down toward the median line of the keel. Feathering on the breast begins at the upper forward parts and progresses down- ward toward the belly. On the neck it progresses from the crown toward the back and from the chin to the breast, the front centerline of the throat being the last to close in. The wing coverts are fully developed in advance of the flight and the body feathers, the upper cov- erts being completed before the lower. The head region is the last to be completed. While two or three pinfeathers may be ob- served in the crown at two weeks of age, down remains in the auricular region and chin until the bird has completed the change to the juvenal plumage over the rest of the body. The first to appear on the head are the elongated crest feathers. The succession progresses from front to back on top of the crown then down toward the eye. The forehead and upper aur- icular region are filled in simultaneously with all feathers appearing at about the same time. As a general rule, it may be said that the succession of feathering progresses in the direc- tion in which the feathers lie. It is well to remember, however, that development begins al- most simultaneously in each of the topographical regions as outlined in the foregoing. By the time the chick is two weeks of age, pinfeathers of the juvenal plumage appear in all of these regions. The replacement of juvenile by adult plumage as well as the annual renewal of feathers in mature birds is followed in the same relative order. While there may be some variation among the individual birds, it follows in general the pattern and succession as that from natal to juvenile. Adult Plumage This is the plumage attained by the mature birds in the fall. If a line or division can he drawn between the end of the juvenile and the beginning of the adult plumage, it might well be established at seven weeks of age, for by this time the tail has attained its growth and the feathering on the head is complete. While down is still present in certain regions notably on the upper thigh, lower sides of the rump, forward of the scapulars and portions of the upper breast, mature plumage is in evidence over most of the body. At this age many pin- feathers can be seen protruding from the skin with the juvenile feathers appended to the tip of the sheath. These pinfeathers may be recognized as the adult plumage by their pigmenta- tion; some being white, others rufous or gray. It was noted that the juveniles remained ap- pended in the same manner as the natal down was appended to them. This was observed in all regions except in the flight feathers and the tail. The adult plumage, however, does not approach completion until the bird attains the age of 18 to 20 weeks, ahhough this is subject to some variation. At the Research Center most of the birds had acquired their mature plumage by October first. But even in the fall the plumage cannot be considered as fully complete since new feathers continue to grow as the season progresses, and probably do so well into mid-winter. As the temperature decreases during the winter months it may be noted that the feathers are much tighter and more closely interlocked than during the fall. At this time all of the spaces are concealed by a thick mat of feathers. By the time s|)ring arrives a slight change in coloration is evident, particularly in the head region, due not to moult but to feather wear. As the season advances, however, certain feathers are moulted about the body. It may be noted that such feathers are dropped from areas ad- jacent to the spaces and a slight thinning of feathers occurs all over the body in general. It I day 1 week 2 ut'eks 3 weeks ^ijV ■"''■V ■/ weeks 6 weeks FIGURE 7. CUOl'SE DEVKI.OPMENT FROM IIM(II1M. TO Mill TIU>n|i B^^:^'t^:^W 8 weeks /O-/:; weeks ^^V) /'A ^'^^ s ',\ I /o-/J weeks a4 THE Sl'ECIES ITS TAXONOMY, RANGE, BIOLUGY, & ECONOMIC IMPORTANCE appears that this is the only period during the course of the year when an actual moult oc- curs which is not followed ininiediatel) h\ feather replacement. Dwight refers to this as the post-nuptial moult. Most feathers which droj) at this time are not renewed until f;di. Il seems logical to assume that su^ With respect to tiic rclalioM>iii|i nf pluniagc dcvclopinciU to the age of tiic ( iiick dining the brood period, an attempt has been made to describe the principal features to be looked for each week from hatching until the bird may be considered adult. Figures P. and ') cor- respond to the age intervals reiognized and show graphically the develoiunenl ol ihc lliiihl feathers which, owing to their regularity of n-placcnicnl. fiirnisli the most conxcniciit iiiilcx for this purpose. * Set- diBcuBsioii ul Analoiiiy, p. 739. GROWTH AND DEVELOPMENT 85 It may be seen that at hatching only seven primaries and nine secondaries are present, although the full complement comjjrises 10 and 15 respectively. For the purpose of this dis- cussion the primaries are numbered with the first at the outermost portion of the wing and the tenth closest the body. Among the secondaries, the short axial feather adjacent to the tenth primary is designated as 1 and the others numbered consecutively toward the body. This order of numbering is most generally followed in ornithological literature although a different system is frequently followed in discussions dealing with domestic fowl. Among the primaries, development progresses in a regular order from number 10 toward number 1, while among the secondaries the pattern is not as regular. The succession here seems to run in the following order: from 3 to 1.5 consecutively with number 2 appearing simultaneously with number 5 and 1 appearing simultaneously with number 6. One Week By the end of the first week considerable growth ma) be observed in the flight feathers. In addition to the seven visible at the time of hatching, numbers 1 and 2 are usually just protrud- ing from the skin. The longest primary of the folded wing extends back to the tail fuzz. In addition, three or four scapulars are present on each side where the wing joins the body. Two Weeks By this time growth of the ju\cnile ])rimaries number 10, 9. and 8 has been completed as evidenced by the absence of blood in the base of the quill where it enters the skin*. The first primary is slightly longer iIkui the overlying greater covert. The axial or number 1 secondary is half grown, while luiiiilx-rs 2 to 1 1 are from two-thirds to one-quarter devel- oped and number 12 is in the |>iii stage. Ten or more scapulars are now apparent on each shoulder and feathers are visible on the sides. Pinfeathers with the downy tufts of the natal |)luinage still appended are present in the region of the lower nape, back, rump, tail, belly, and (lank. Two or three pinfeathers. which later develop into the crest, may be discerned on the crown. The lower wing coverts are still obscured bv down. Three Weeks The juvenile primaries number 10 and 9 have already dropped out and are being re- placed by adult feathers. The tip of number 10 is exposed and slightly longer than the base of the quill. Number 9 cither may be missing or the tip barely showing through the skin. Of the juvenile flight feathers still present, growth is complete in primaries number 8, 7 and 6, as is also true of secondaries number 3 and 4. The greater wing coverts are fully developed and the upper surface of the wing is well cov- ered except in the area of the web. This latter region is sparsely covered with down with no pin feathers in evidence. The edge of the web. however, is covered with three rows of feath- ers, growth in the first row being complete. The up])er part of the head is completely covered with pinfeathers from the base of the mandible back to the nape. That portion below the eyes and under the chin is still in down. Juvenile feathers have appeared on the back of the neck merging with the scapulars. The complete bridge of feathers across the back has been acquired at this age. The feathers on the sides have attained sufficient length to cover a portion of the belly. This latter re- * After growth is cnniplele a t)iiill becomes transhireiil and dry. 3 6£ ns a V JO u o u in D O fiS c o z. 3 o z u CL. o > Id Q OS Id SB •< b) (K o z O z o Id OS O O Bfi CL. Id OS 3 O 88 THE SPECIES— ITS TAXONOMY, RANGE. BIOLOGY. & ECONOMIC IMPORTANCE gion is still mostly obscured by down and no pinfeathers appear on the median line of the keel. Pinfeathers with downy tufts attached cover the back and rump from the neck to the oil gland. These feathers have attained sufficient growth so that they are partially visible through the downy fuzz. On the rump the tips of a few feathers are showing beyond their sheaths. While the outer tail feathers are still in the pin stage, the central ones have emerged from their sheathes. Four Ifeeks At this age adult primary number 10 is about half grown, number 9 is about one-third de- veloped and number 8 is in the pin stage. Growth is complete in juveniles 7, 6. 5 and 4. Among the secondaries the axial as well as numbers 2 to 10 inclusive are fully developed while number 15 is well out of its sheath. The body of the bird in general appears well feathered in juvenile plumage. The side of the head still appears downy. The pinfeathers on the chin also are obscured by down. The juvenile feathers on the sides of the neck cover the downy area of the front median line of the throat. The same is true of the feathers which cover the downy median line of the belly. Numerous feathers are noted on the legs with those on the upper tarsus haxiiig downy tufts appended. Five Weeks During the fifth week, although none complete their development, growth progresses rapid- ly among the adult primarv feathers numbered 10. 9 and 8. Number 10 is now approxi- mately the same length as the lunnber 2 juvenile secondary. Juvenile primar\ number 7, how- ever, has been dropped and the tip of the adult feather is exposed. The bird now has a sleek appearance, having lost the fuzziness due to the many appendant downy tufts which were prevalent during previous stages of development. Growth is complete in many of the body feathers. The tail, however, is still growing. Six Weeks Among the flight feathers adult primaries number 10 and 9 have completed their growth and number 6 has appeared as a pinfeather. Juveniles 5, 4 and 3 have also become com- pletely developed. Among the juvenile secondaries, number 11 has completed its growth. It should be noted particularly that two of the adult primaries have completed their develop- ment before the juvenile primaries numbered 1 and 2 are half grown. The juvenile tail has completed its growth. Seven Weeks Though the flight feathers continue their regular growth, ihc only significant change in feather replacement is the juvenile secondary number 12 which has completed its growth by the end of the seventh week. Over the body in general the juvenile plumage is now com- plete, with niost of the feathers very loosely htld in the follicles and released readily when the bird i,~ handled. Evidence of the adult body j)lumage begins to appear in the form of pinfeathers according to the regular succession outlined in the discussion of juvenile plumage. Some spaces and areas adjacent to them are still comparatively bare with down remaining in most cases. GROWTH AND DEVELOPMENT 89 Eight Weeks Among the adult primaries in addition to number 10 and 9, 8 is now fully developed. Numbers 7, 6 and 5 continue their growth. Growth is complete in juvenile secondaries from number 9 to 15 and the adult feathers from number 1 to 8 are developing^ numbers 1 and 8 being still in the pin stage. The back and rump are completely covered with pinfeathers of the adult plumage. In practically all cases the juvenile feathers are appended to the tips of the sheaths. This ar- rangement causes the old juvenile feathers to be lengthened and thereby protects the new feathers coming in under them. The natal down which is still present in areas previously described is being forced out by adult pinfeathers. In these cases the intermediate or juvenile plumage is omitted. The downy tuft is appended to the tip of the aduh pinfeather. Nine Weeks Juvenile primary number 4 has been dropped and is being replaced by the adult primary. In like manner numbers 9 and 10 in the juvenile secondaries have been dropped and are being replaced by adult feathers. The adult bod) i>lumage continues its growth at approximately the same rate as the juve- nile developed previously. The general pattern fallowed closely |)arallels the succession of feathering from the third to the eighth week. Ten Weeks Growth of the adult flight feathers continues with only the number 11 juvenile second- ary being replaced by the adult feather. The growth of the number () adult primary has been completed. Numbers 1 and 2 [)rimaries have nearly completed lluir development. Juvenile tail feathers drop out between the ninth aiiT AND 108 YOUNG GROUSE COLLECTED IN NEW YORK ST VTE— 1931-1911 Number specimens Average weight in grams Average wei ;ht in ounces Male Female Male Female Male Female Adult June 27 43 26 i3 7 9 11 13 15 7 6 33 51 to 3 8 6 13 12 15 15 10 8 615.0 607.3 604.0 635.6 654.1 650.7 616.9 636.9 (>15.5 (>25. 1 f>21.7 .500.1 .508.1 517.1 523.3 573.3 576.8 586.8 579.3 549.4 518.3 5U.8 5.36.6 21.9 21.7 21.5 22^7 23.3 23.2 23.1 22.7 21.9 22.3 22.3 17.9 July 18.1 August 18.5 S:!ptembcr October November December .lanuary February 18.7 20.1 20.6 20.9 20.7 19.6 18.5 April 19. t 19.1 Young September October 13 15 10 8 ' \ 5 \ 17 6 •1 7 560.2 .591.5 .597.2 619.1 612.8 597.4 450.2 187.8 .521.5 529.0 5 16.6 520.6 511.4 20.0 21.2 21.3 22.1 2L8 21.3 16.1 17.1 November December 18.7 18.9 19.5 February March 18.6 18.2 Seasonal Trend An analysis of weight trends and differentiations as here illustrated (figure 10) raises many an interesting question of influence and relationship. In distinct contrast with the Scotch grouse of Great Britain, ruffed grouse of all ages and sexes in general seem to follow much the same trend at approximately the same time. They reach their highest weight of the year in November or December following a period of fall plenty. The average weight of New York grouse at this season is about 1 lb. 7 oz. (651 grams) for adult males and 1 lb. 5 oz. (587 grams) for adult females. With the onset of real winter, certainly by the middle of January in New York, a loss in weight sets in, gradual at first but more rapid as the winter progresses. This culminates about the middle of March with an average loss of from 1^,4 to 21/^ oz. (35 to 69 grams). y2 THE ^I'ECIES^ITS TAXU.\ 0,1/1 . RA.\GE. BIOLOGY. & ECO.\ OMIC IMPORTANCE Over the next month, the birds regain some % oz. 1 9 to 27 prams) of the weight they have lost. Possibly the melting snows by then have uncovered foods of greater nutriti\e value or, perhaps with spring's warmer weather, the energy outgo is less than that provided by the food taken in. At all events, the increase is short-lived for, through the active days of May and June, more weight is lost. The low point of the year, for females, is about June 15. for males about August 15. when they average about 1 lb. 2 oz. I 500 grams) and 1 lb. 6 oz. I 604 grams) respectively. The demanding responsibilities of courtship, of starting a family, and of moulting about over, their weight rises, slowly at first, then more rapidly throughout the early fall to its November peak. oz 24 CMS , eao 1- I 20 ' / / (J 1 S60 ' / ' 1 / Moles- Adult -— - Males- Young ^^ Ferrlales - Adul+ — — Femal es - Young SEP OCT. NOV DEC JAN FEB MAR APR MAY JUN. JUL AUG SEP 15 15 15 15 15 15 15 15 15 15 15 15 15 MONTHS FIGURE 10. AVERAGE SEASONAL WEI(;HTS OF 394 ADULT AND lOii YOUNG WILD GROUSE FROM NEW YORK Differences Between the Sexes Many a biologist, in attempting to identify the sex of grouse in the field, has been puzzled by the over-lapping of indicative characteristics*. Among the best of these is liic length of the tail, for in males it will a\crage nearb three-quarters of an inch longer. By study- ing the weight graph, one also finds that thi> rule-of-thumb judgment of many hunters by which the heavier grouse are considered to be males, iiolds good in gener;il. During the hunting season, the male adult will usualK average aliout 2-3 oz. (77 grams I lii'a\ icr tliaii its mate. Even a male of the year, while a\eraging about 2 oz. (57 grams) lighter than one of the preceding year, is, nevertheless, apt to be some % oz. (20 grams) heavier than an adult female. Weight must, of course, be accepted merelv as a general indication of sex. for many birds of each group are killed which are above or below the levels here given. Usually, however, a heavy, long-bodied bird is a male; a plum]), fairly light bird, a female. On the average. an adult female is still 2/3 oz. (17 grams) liglilcr. i-vcn at her December peak, than is the * Sep di»cu»Bion iindrr Hnw to Krroiinilc Sex, p, 39. WEIGHT 93 male at his lowest weight in August. It is at year's end that the weights of the two sexes most nearly match, for the birds are then only about 2V4 oz. (64 gramsj apart. Curiously enough, the weight difference is greatest early in June shortly after the strain of egg laying and subsequent incubation is passed. The female loses rapidly during the nesting period, reaching the low point, about 4 oz. 1.115 grams) below that of her more carefree mate, shortly after the chicks are hatched. It is not always easy to ascribe reasons for the variations in the weight pattern of either sex. The influence of good food and of freedom from reproductive activities offer likely ex- planations for the rapid gain of weight in the fall. Through the winter, the female appar- ently loses weight more rapidly than the male but, like the Scotch grouse, with the coming of spring, some is regained. Losses during e^g laying (mainly mid-April to early May in New York) are not large. But the subsequent confinement of incubation brings on a swift and deep decline in female weights, possibly in response to restriction of her opportunity to feed freel). Al the low point, ihc average female will weigh but 1 lb. 2 oz. (500 grams). This is probably dangerously low, for then it is, in raising grouse in cai)tivity, thai they are most likely to die from chronic diseases. Throughout late June and July, the male is mouUing and slowly losing weight. Yet the female, though likewise renewing her feathers, gains steadily. Grouse are able to call rather quickly on their chemical reserve once their metabolic reserve has been exhausted. It is prob- able that the hen approaches this condition earlv in June, the result of having already lost almost one-sixth of her December weight and that from this time on a slow recovery takes place. This may conlrii)Ulc Id the aliilitv of the female to i)ul on weight even though moulting and caring for her brood. During the latter half of September or early October, she gains almost 2 oz. (about 50 grams! to put her once more in good flesh. The male neither varies so greatly in his weight throughout the year, nor are the changes as rapid. His drumming and strutting, so prevalent in early spring, do not prevent him from gaining slightly as winter wanes, though the increase is not as pronounced as with the female. By early June, though liis breeding dis|ilay is now largely an activ ity of the past and he has scarcely begun to moult, his weight, curiously enough, is still dropping. This continues slowly throughout the period of feather replacement and well into August. In the fall, he apparently requires some three months to put on the same amount of flesh that the female acquires in one. <) I THE SPECIES ITS T.iXOXOMY. RA.\GE. BIOLOC). & ECOXOMIC I MI'OKTANCE Agk Differences Pvormally, a grouse chick at birth weighs nearly !/•> oz. (] 1 to 13 grams), though this varies slightly from year to year. Because of the difficulties involved, few youngsters have been banded and subsequently recaptured and weighed in the wild. Weekly weights of birds raised in captivity, however, have been kept (table 11). Comparing these data with those in figure 10 it will be noted that while the pen-reared females attained somewhat higher weights by early fall than the average for birds collected in the field, the reverse was true among the males. From September's frosty nights, the birds of the year seem to follow much the same weight pattern as do their elders (figure 10). Although they are then some 2y-j, oz. (60 to 70 grams) lighter on the average, they are probably even more active foragers, for they rapidly nar- row the gap. Too, the gain is continued later, for apparently they do not reach their peak until sometime in January, although they still average upwards of 1 oz. (20 to 30 grams) less at this time. Also, birds of the year seem to lose a little less throughout the rest of the winter, though they are still some 8 to 16 grams lighter as late as the first of March, the greater difference being recorded among the males. Beyond this time they usually be- come indistinguishable from older birds, due to the disappearance of the bursa and the moult- ing of their two juvenile outer primaries. TABLE 11. AVERAGE WEEKLY WEIGHTS. IN GRAMS, OF 8 MALE AND 7 FEMALE HAND-REARED GROUSE CHICKS Sen Weight at hatching Weight at weekly intervals after hatching 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Male 12 21 39 64 95 132 180 237 295 350 400 442 479 503 525 540 550 559 567 571 579 Female 11 20 32 53 81 119 163 213 260 309 352 394 428 452 470 484 498 502 508 510 Sll By the middle of September, the weight difference between the sexes among these younger birds has apparently reached its maximum, for the female is nearly 4 oz. (110 grams) the lighter of the two, the average being 1 lb. (450 grams) and 1 lb. 4 oz. (560 grams) re- spectively. Even at their January peak there is a difference of close to 3 oz. (75 grams) between them, a spread, however, not much greater than that between their elders at this point. Differences in weights of grouse shot during the hunting season may often be ascribed to age differentials for even then the birds of the year average 2 oz. less in weight than do old birds of the same sex. Kegional and Yearly Differences To secure an accurate picture of regional and ycuil) differences, it is necessary to analyze the weights of an adequate number of individuals collected from each region each year. Un- fortunately, full grown birds collected prior to 1939 were not separated into adults and birds of the year. At that time the reasonably accurate means of identifying the latter by the presence of the bursa of Fabricii or an examination of the tips of the two outer primaries, had not been recognized. A careful analysis of the resultant weight data for each age group for 1939-41 reveals no significant differences cither between the three major regions of the WEIGHT 95 state or between years for the state as a whole during this period. From 1936 through 1940, Dr. A. L. RomanoS conducted for the Investigation a detailed study of annual variations in the development of the embryo*. Slight weight differences year by year among the eggs and among the chicks upon hatching were recorded. Their signifi- cance, however, is not yet clear. It is to an expansion of this study of the causes of these less obvious changes in grouse weights year by year as well as to other physiological research that one must look for indica- tions of the part played by food and, indirectly, by weather, in influencing periodic decreases in grouse abundance. Many more eggs and birds will have to be collected over a much wider area, and better techniques for their physical and chemical examination worked out, before the indications that surely exist here of the relationships of these environmental factors to cycles can be adequately outlined or understood. Extremes of Grouse Weight Few subjects provoke better arguments among grouse hunters tiian does the weight of the largest grouse ever shot. Nearly every year, birds are killed weighing over 1% lb. Pubhshed records on this subject are numerous. "Sus Q. Hannah'"^, who handled many grouse for the markets, claims that the weight "varies from year to year according to circumstances, food, weather, etc." In 1930, there passed through his hands a cock grouse that is said to have weighed 30 oz., even though no unusual amount of food was in its crop. Writing in "Forest and Stream", in 1880, Fellows'" cites a bird killed near Hornellsville, l\. Y., which weighed 321/2 oz. A still larger bird which "weighed tliirty-three ounces on tested scales" has been reported"'" as having been killed many years ago in Rutland, Mass. But what seems to be the heaviest bird reliably recorded was weighed by John Burnham'"'. Shot in Essex County, N. Y., in the Adirondacks, it tipped the scales at 2 lb. 4 oz. The Investigation has not weighed one which has approached this Goliath. Males usually weigh more than females and birds two to three years of age more than birds of the year. Probably therefore, all the above records are of old males. In the other direction, adults may be reduced to skin and bones and still cling to the breath of life. One female, found in December 1940 in Chenango County, N. Y., and suf- fering from chronic stomach worm infection, weighed but lO'/a oz. (292 grams). Altogether too weak to fly, it was easily caught by hand and died soon after. Weight as an Indicator of Health Weight is still the best easily available measure of grouse health. Its fluctuations are enough, however, to be confusing. Weight, as an index of condition, must, therefore, be compared with the normal variation encountered among healthy individuals month by month. This differs seasonally as well as among individual birds during one season. But in general, there is a zone of good health (figure 11) extending above and below the average within which variations in the weights of most healthy individuals may be expected to fall. Below this is a danger zone. Weights falling here are apt to indicate environmental difiicuhies or the presence of serious disease. Beyond this danger zone is a critical or lethal zone. Birds whose weights fall within this may seldom be expected to live for long. It is recognized, however, that there exists a point above the weight at death below which, due to metabolic and chemical exhaustion, the system will not recover even though the bird * See discussion of Embryology, p. 72. 96 THE SPECIES ITS TAXONOMY, RANGE. BIOLOGY, & ECO\ OMIC IMPORTANCE OZ. CMS. Males 25 |1 710 24 K 680 23 J 650 22 \ 620 21 J 590 o u 5 20 19 18 560 530 1 500 17 J 470 16 1 440 15 J 410 14 1 380 OZ CMS 22 K 620 21 J 590 20 \ 560 X UJ 19 18 j 530 500 $ 17 16 15 14 1 470 440 410 380 JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC JAN 15 15 15 15 15 15 15 15 15 15 15 15 15 MONTHS Femoles JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC JAN 15 15 15 15 15 15 15 15 15 15 15 15 15 MONTHS ® Average weight by months. ® Approximate lowest weight for good health. © Approximate critical weight. @ AVERAGE WEIGHT AT DEATH. IK. IKK II. SK.ASONM. U IK.IIl -11 KAl.TIl UKl. \'H((\.s| 1 1 p o|. \I)U.I (.Hoist I I Zone of Good Health I I Danger Zone Critical Zone WEIGHT 97 may still live for days or weeks. Naturally, this critical point may be expected to vary somewhat with the environment and the individual. Though known for the ring-necked pheasant in captivity, no exact measurement, covering a sufficient number of specimens, has ever been made to secure a reliable average for grouse. Long*, however, has worked out for the Investigation an approximation for adult hand-raised birds at the peak of their weight cycle. Roughly the line falls between about 16 oz. (460 grams) and 14 oz. 1 390 grams) in the case of males and for females it lies between about 14 oz. (400 grams 1 and 12 oz. (350 grams). It represents about a 30 per cent loss from original weight among both sexes. GROUSE AT THE KESEARCH CENTER WERE WEIGHED PERIODICALLY AS ONE MEANS OF COMBAT- TING DISEASE Further light on what constitutes the danger zone and the critical weight is gained by analyzing weight losses under various conditions, such as extremes of temperature, lack of food, intense activity and niouhing. One indication of the threshold of the danger zone is the point at which birds are likely to fall prey to chronic diseases. At its lower limit are the weights under which a bird cannot drop and live for long. This has been roughly deter- mined for New York grouse by a study of the final weight decline of birds which have died of chronic ailments. Nevertheless, this conclusion must be interpreted as only an estimate, * See discussion iiiulcr Pliysinlugy. p. 60. 'j;i THE SI'KCIES ITS TAXONOMY, RANGE, BIOLOGY, ct ECONOMIC IMI'OKTANCE carefully based on the meager facts available. Perhaps the clearest idea of the weipht-health relationship may be gained by studying fig- ure 11. By comparing the weight of the birds collected or picked up dead at any period of the year, with the average here presented, a fair diagnosis of their relative state of health may be made. Of course, one may also utilize his own judgment of the condition of the bird, but unless one is thoroughly skilled in such matters, rule-of-thumb methods, such as the mere hefting of a bird shot in the field or estimating its condition by the relative sharpness of the breastbone, are notoriously unreliable. Many times grouse have been shot and sent to the Laboratory, as seriously diseased, simply because a few of the long roundworms ( Ascaridia) , found upon cleaning, have predisposed a hunter to judge that the bird was poor in flesh. Actual weight comparisons with those given in the table, while of course in no way absolute, will provide, in most cases, the best method of appraising in New York, and probably throughout the Northeast, the general health of grouse. MEASUREMENTS In general, adult grouse do not vary greatly in size, especially with respect to their skeletal structure, although males tend to be slightly larger than females. Unfortunately, time has not permitted detailed measurement of the majority of the specimens which the Investigation has taken. With respect to the eastern races of the species, the following as recorded by Forbush'", is representative. "Length 15.50 to 19.00 in.; spread 22.00 to 25.00; folded wing 6.90 to 7.50; tail 5.50 to 7.00; bill from nostril .50 to .53; tarsus 1.50 to 1.60." An interesting note by Poole™ gives 527 sq. cm. as the wing area of a grouse which weighed 516.5 grams, or a relationship of 1.02 sq. cm. per gram. In a series of adult birds collected in New York State between January and April, however, a record was kept of the length of the middle tail feathers, the results of which follow. TABLE 12. LENGTH OF CENTRAL TAIL FEATHKHS OF ADULT GROUSE FROM NEW YORK STATE Number of specimens Average Longest Shortest Male Female 31 20 6.56 in. 5.43 in. 7.56 in. 5.75 in. 5.87 in. 5.00 in. It should be notcli of the bird is liighK palalalije when properly cooked. 1 hat the Indians probably made considerable use of it. especially during its ])eriods of abundance, was indicated bv Hanm de I.ahontoii"'' ^vh^ reported in 1703 that tliev "shot th<'m with ar- Hy Ganliiirr Hump. ECOIKOMIC IMPORTANCE 101 rows, for they were not worth a charge of powder". From a quaint tract entitled "'New England's Plantations""", published in 1630, one gathers that turkeys, grouse, geese and ducks played a major part in the daily dinners of the earliest settlers. Utilized at first for personal use only, as time went on grouse became increasingly popular as a commercial food. Two score years ago it required hundreds of thousands of grouse an- nually to satisfy the hungry markets in the larger cities. Prices, where birds were scarce, ran as high as $5.00 or even $7.00 a pair, though during periods of abundance they ofttimes sold for 50 cents apiece.* With the prohibition of its sale, grouse have come to be looked upon more as an excuse for a keen day's sport than merely as an article of food, even though today they remain one of the greatest delicacies of the hunt. The best figures available indicate that upwards of two million grouse may be shot annually throughout its range during periods of normal abundance. The average bird will weigh about a pound and a half in the field. Properly prepared, the annual harvest may furnish upwards to four niillii>n meals. The Importance of Food Preferences Though grouse are highly insectivorous as youngsters and later great seed-eaters, they can- not compare with either the ring-necked pheasant or the bolnvhite quail to say nothing of many of the smaller birds in their consumption of items detrimental to agriculture. This is only natural, since good grouse coverts are never highly cultivated. Their food consists largely of woodland fruits, berries, buds, seeds and leaves, with minor emphasis on insects in the summer. For the most part, their feeding habits neither citnllict with, nor contribute to, man's interest. True, some seeds are not digested and many wildlife food |)lants are thereby more widely dispersed. But it is mainly among the insects that the grouse makes its relati\ely small con- tribution. The list of species picked up. largely by the chicks, includes, according to McAtee and Beal ' ', the Colorado potato beetle, clover root weevil, pale strijied flea beetle. grape\ ine leaf beetle, red humped apple caterpillar, grasshoppers, cottonworms. army worms, cutworms, sawfly larvae and May beetles. To this group our own studies have added adult sawflies, the strawberry weevil, black vine weevil, poplar borer, cucumber beetle, elm leaf beetle, maple- tree worm, canker worm, and various plant bugs and leaf hoppers. Though ants, beetles, and caterpilla-s are taken primarily among this group, the great abundance of insects in general'^, as well as of other foods, and the relatively large territory available to each brood, makes it doubtful if grouse alone, even where they are plentiful, make much of an impression on the numbers of harmful species. Nevertheless they do. of course, contribute a share to the aggregate beneficial effect of birds in holding in check the hexapod hordes. * See ChaptiT I. p. 8. A Sporial stiiilirs ..I iiis.-it al.iin.lan.-.- in Jim.' .it IIM an.l 1937 nn llio Connrrlicul Hill an.! A.lirnnilack areas rt-vcaled an average of over 300.000 indiviiliials per aere in good grouse cover (Biimmer feedinK Eroiindl. 1(12 THE SPECIES— ITS TAXONOMY, RANGE, BIOLOGY, & ECONOMIC IMPORTANCE Budding of Fruit Trees At first thought, it seems rather incongruous to present-day observers that the grouse may at times be an economic liability. Nevertheless, as early as 1870, certain Massachusetts towns instituted a bounty of 25 cents on this species because of its alleged damage to apple orchards and fruit trees in general through eating the buds during the winter and early spring. In Kentucky also, according to Forbush"* they early acquired a reputation blacker than that of the crow, not only for their apple budding propensities, but also because they were said to pick up sown grain from the open fields — a supposition not reported in any other part of their range. Apparently, however, New England grouse are the chief offenders, for among these rugged hills apple trees have been planted invitingly close to good grouse coverts. Usually but three or four birds are involved at any one place, but there is one report"" of eighteen birds simul- taneously working on one tree. Though food equally attractive to most grouse be abundant close by, a few birds thus concentrating on an orchard may easily strip many a limb almost bare of buds. Maynard^ mentions a grouse he shot about ten o'clock in the morning from whose crop he extracted 180 apple buds, although the average number taken is apparently much smaller. Opinions as to the effects of such depredations vary with the interest of the individual and the severity of the pruning experienced. Complaints usually are limited to periods of grouse abundance and are handled in New York State by giving the harried landowner permission to shoot the culprits. But other more strenuous measures, such as the Massachusetts bountv, have occasionally been resorted to elsewhere. Probably the most paradoxical situation is to be found in the record of the State of New Hampshire, where a statute was passed in 1915 requiring the state "to pay for any damage to 'annual crops or fruit trees by game birds or game quadrupeds protected by law' ", such payments to be made "out of fish and game funds, which accrue entirelv from license fees, fines and forfeitures"^'. Under it, claims at first were few, largely because the law was little known and grouse were going through the periodic scarcity of 1917-18. Then the abundance of birds surged upwards and. with it, damage claims. It is even said that the Farm Bureau Federation and the State Horticultural Society started membership campaigns, using as a lever the assistance they might offer in settling complaints. By 1923, the situation had be- come so wides])read that over 400 such claims were presented for adjustment. Since July 1. 1922, New Hampshire has paid approximately $70,000* for damage by grouse in spite of an investigation being made in each case. The outstanding years (fiscal), by far, were those ending June 30. 1924 and June 30, 1927. when $26,800.72 and S13.989..S0 were paid re- spectively, the former amounting to almost one-quarter of the total income of the Fish and Game Department. But even in the years 1927-28 and 1928-29 when, to many observers, the future of the grouse over much of the Northeast seemed in danger, payments of $4,152.21 and $1,047.60 respectively, were made. In 1941, over $1,000 was paid. But, even as the controversy raged, many individuals were of the opinion that grouse bud- ding results in no appreciable damage to the tree. As a rule, the birds confine their activities lo the top and renter portions. Thus it often happens thai trees, which have been quite heavily i)udded, produce on their sides and lower braixhes fniil of a ])etler si/e and qnalily than those which have not been so pruned. Carprnirr, R, C |irr§niial ii-tliT In llir ntithnr,. Dri-riiibrr 2ti 1111. gguw^TMW*^ ,. (tie Cmtrnj/MX. W— - -?C ECONOMIC IMPORTANCE 103 Whether this be an advantage to the orchardist or not depends upon conditions. Thus Judd"^ cites one case of an apple tree pruned severely two years in succession, yet produc- ing "barrels of apples, all nearly perfect and fair, just the result of a vigorous trimming." On the other hand, Bartlett'^, described an orchard planted on fertile land and well cared for throughout, one side of which adjoined inhabited grouse cover. The production record for both portions was about the same until the birds discovered the apple buds in the block closest to their covert. The result is described in these words: "In the spring the broken ends of limbs and white spots where buds had been taken were so numerous that the trees looked white as you approached this section of the orchard. As you came nearer casual observation would almost make you believe that no fruit would set. and that very few leaves would appear. At blossom time, flowers were few and scattered. At fruit picking time, these trees had apples of very good quality, but not 50 per cent of the amount of fruit in bushels borne by the other sections of the orchard which had not been budded. The owner set the shortage of fruit at 200 barrels." One must bear in mind, however, that the latter represents the exception rather than the rule, considering grouse range in general. And today, throughout the Northeast, one finds abandoned many farms from which damage complaints once originated. Then. too. hunters in increasing numbers, dissatisfied with the slower ring-necked pheasant, are again turning their attention to grouse. One has but to advertise a concentration of birds sufficient to cause damage, to interest local sportsmen in assisting in their control. Grouse as a Game Crop With the outlawing of market hunting, the shooting of grouse became a sport to be indulged in primarily for recreation. Furthermore, the harvesting of the crop each year has con- tributed in many ways to the revenue of many states and the income of their citizens. Tt has been estimated that the justly famous red grouse moors of Scotland and England I>rought in about 1,300,000 pounds ($6..'500.000) in gross rent per year before World War TI. In Scotland the "glorious 12th" (August) which commences the shooting season is a sporting and social event of the first rank. The railroads run special "grouse trains" and the day is celebrated in song and story. On these moors was provided shooting of a kind largely un- known in this country, since it was not unusual for a relatively few shooters to kill in excess of 6000 birds in a good year on a sinp;le properly managed moor. Over here, ducks, pheasants and Hungarian partridges occasionally reach concentrations which miciht permit bags of simi- lar size, though largely on private shooting preserves. But the ruffed grouse is not particu- larly gregarious and though it has been known to reach fall concentrations as high as one bird per two and a half acres, in individual coverts, efforts to maintain such abundance have not been successful. Yet. for its sporting attributes, the bird is particularly esteemed among a large group of sportsmen who glory in the difficulties of the hunt. It is widely distributed and occupies for the most part land on which no hunting restrictions have been placed. These factors combine in New York alone to put tenfold more nimrods in the field in its pursuit than take gun to himt the red grouse throughout the entire British Isles. No figures as to the number of grouse actually killed in any one of the several states are available today nor is the number of hunters seeking the species accurately known. In fact, but twelve states have even set up the machinery to gather aught save the most general of estimates on their game take. Even in New York, where the law requires such records of every licensee and all are tabulated each year, the actual number of grouse taken is substan- tially in excess of that reported. In a normal year, when sportsmen's license returns indicate 104 THE SPECIES—ITS TAXONOMY. RASGE. BIOLOGY. & ECONOMIC IMPORTANCE the take to be about 170,000 birds, it is safe to say that the actual miiiihcr sliot would ap- proximate 300,000. Field studies conducted by the Investigation in certain Southern Tier ( N. Y. ) counties over two successive years, indicate that about 17 per cent of the fall pop- ulation of adult grouse in that region are killed by hunters in a season of average abundance. A take (jf 300.000 birds, then, would indicate a p<)|)ulation of approximately a million and three-quarters in i\ew York alone. From an inspection of license records, one finds three to be the number of grouse reported most frequently taken by individual \ew York State hunters. Yet the number is undoubtedly lower in actuality, for most grouse here are so hard hunted and wary that some grouse hunters are entirely unsuccessful and thus make no report. Rut, if one accepts three as the average kill it follows that there are upwards of 10(1.000 grouse hunters in this State alone. Over its entire range, based on a canvas of existing records and estimates, a reasonable ap- proximation of the annual kill during a year of average grouse abundance, would amount to at least 1.900.000 birds. If, by way of a very rough inference, one were to apply New York's kill ratio to this, an estimated total population of 12,600.000 would result. Yet it is well known that these birds are tamer and, therefore, easier to kill in many parts of their range than they are in the Northeast. To compensate for this the hunting pressure is probably less. Thus, should grouse hunters over the whole area occupied bv the species aver- age four birds each per season, one might conjecture that there are perhaps 500,000 sportsmen who make grouse hunting their hobby. Those who know New York grouse will agree that the estimates presented for that section well reflect conditions as thev exist. In perhaps six or eight f)ther states the figures covering the annual kill of grouse are equally indicative. For the rest it has been necessary to fall back on the ofttimes unreliable method of "best estimates". But. in spite of the obvious inaccu- racies which beset such efforts, the picture is significant. The ruffed grouse, unlike its Scotch cousin, represents a crop, the size of which is still controlled largely by the whims of nature. Yet it furnishes recreation to perhaps half a million hunters instead of the comparatively few who are able to enjoy this sport in the British Isles. Furthermore, the actual revemie derived from hanesting the crop is appreciable. An expenditure by each grouse hunter of only about 816.00 a year for guns, ammunition, hunting clothes, travel and lodging would roughly equal Great Britain's revenue from the rental of its grouse moors. Sai.k of Breeding Stock Grouse for restocking depleted coverts have brought prices well ;ib(i\c those paid fur most native game birds for many vears. Though current prices (19-12) for wild trapped birds are S6.00 to SIO.OO apiece, they ha\e run both above and below these figures. l'tMha|)s 1.(^00 birds have been tra|)ped. mostlv in Canada, transported and released to restock dciilrtcd coverts and bring in new blood -an idea widely, though probably erroneously, accepted as necessary. Hand-raised birds, too. are at a ])remium. for there are but one or two places where breeders have succeeded in establishing a strain that will do more than reproduce itself in captivity without frequent additions of birds or eggs from the wild. Such hand-raised birds have brought prices of from $10.00 to S.SO.OO a pair, with but few birds for sale. Artificial grouse propagation, at least in its experimental stages, is costly and the present supply of hand- raised breeders is. therefore, largely concentrated in the hands of a few jiublic agencies, not- ably New York State. PART II The Factors That Affect Abundance CHAPTER III COVER CHARACTERISTICS AND SHELTER REQUIREMENTS By Gardiner Bump CHARACTERISTICS OF PRODUCTIVE GROUSE COVER In General — Covert Size and Shape — Type Size and Shape — Type Composition — Inter- spersion of Types — By Regions in New York State — The Adirondack Region — The Catskill Region — Rest of State. THE ROLE OF SUCCESSION COVER TYPES RECOGNIZED COVER REQUIREMENTS OF THE GROUSE Nesting Cover — Types of Cover Chosen — Density of Undergrowth at Nest Site — Sites Preferred as Nest Locations — Nest Location in Relation to Degree of Slope — Nest Loca- tion in Relation to Aspect of Slope — Influence of Conifers on Nest Location — Effect of Openings on Nest Location — Effect of Slashings on Nest Location — Influence of Other Types of Openings on Nest Location — Effect of Crown Cover on Nest Survival — Effect of Undergrowth Density on Nest Survival — Thickets versus Open Cover as Nest Locations — Effect of Distance from an Opening on Nesting Success. Brood Cover and Related Influences — ^Types of Cover Used by Grouse Broods — Change in Cover Use as the Broods Develop — Effect of Crown Cover Density on Brood Choice — Types of Cover LIsed by Grouse Broods at Various Times of Day — Choice of Crown Cover as Affected by Climatic and Ground Conditions — temperature — ivmd — atmospheric coridilions — ground conditions — Some Relationships of Undergro^vth to Grouse Broods — Types of Undergrowth Chosen by Grouse Broods at Various Ages- Effect of Density of Undergrowth on Brood Cover Choice — Effect of Miscellaneous Con- ditions on the Types of Undergrowth Chosen by the Brood — L^se of Different Degrees of Slope at Various Ages — Slopes Used by Grouse Broods at Different Times of Day — Some Relationships of Temperature, Wind and Atmospheric Conditions to Slope Use. Adult Grouse Cover and Related Influences — Types of Cover Used by Adult Grouse — Seasonal and Monthly Cover Preferences — Types of Cover Used by Adult Grouse at Various Times of Day — winter — spring — summer — fall — Effect of Various Weather Con- ditions on the Choice of Cover by Adult Grouse — temperature — wind — atmospheric con- ditions— Effect of Various Ground Conditions on the Choice of Cover by Adult Grouse — Preference of Adult Grouse for Ground Over Tree — Influence of Slope on Adult Grouse Distribution — Influences Affecting the Distance Grouse Flush from an Observer. 106 COl'ER CHARACTERISTICS A.\D SHELTER REQUIREMENTS Reflushes — Cover chosen in which to alight — Distance between Flush and Reflush — Effect of Sex on Flight Distance — Tendency to Alight in a Tree. THE ROLE OF COVER COMPOSITION AND ARRANGEMENT THE VALUE OF EDGES Effect on Adult Grouse Distribution — Effect on Brood Distribution — Relation of Edges to Nest Location — Variation in Importance of Edges. THE VALUE OF CLEAR-CUT AREAS ^ SUMMARY No factor is more important than cover in determining grouse distribution and in influencing the productivity of grouse populations, (p. 109). Because of the sedentary nature of the birds, a good grouse habitat must meet all food and shelter requirements of the species within a relatively small area. (p. 110). Grouse cover is constantly changing as one type grows into the next more advanced stage. (p. 118). The outstanding grouse requirements which must be met by cover are for spring nesting grounds, summer feeding areas, fall feeding grounds and winter shelter. (p. 111). Grouse cover is as much a reflection of the various farm and forest uses to which the land has been put as the result of varying climatic, physiographic and biotic influences. (p. 114-11!!). The extent to which such crops as forage or wood products are harvested may affect the pro- ductivity of a woodland for grouse quite as much as does its natural composition.! p. 118). The partridge finds its year-round requirements met neither in the first stages of plant succes- sion associated with grasslands nor in the last or climax types of dense unbroken forest, (p. 118). Some of the most pnidiictive grouse habitats in the State are to be found where woodlands are broken u|) by farm clearings which have been abandoned and are beginning to revert, (p. 116). Nesting cover and site exhibit certain characteristics as follows: Woodland types are strongly favored as nesting cover. ( |). 128). Of these, nests are most likely to be found in second-^Towlh hardwoods; least likely to be placed among conifers, (p. 128). Most birds are reluctant to nest where the undergrowth is dense, (p. 128). Grouse prefer to nest at the base of a tree. I p. 130). Slope, degree or aspect, does not materiaijv iridiicin c llic rliDJcc nf a nesting site, (p. 130). The desirabilitv of a nesting site varies in\ersel\ with its distance from an opening, (p. 134).' SUMMARY 107 Nests in woodland types are commonly located within 100 feet of a woods road, trail or an open field, (p. 135). Nests are most likely to be broken up by predators when in coniferous cover; least likely, if in spot-lumbered areas, (p. 135). About the same number of nests are destroyed by predators irrespective of the density of the undergrowth in the cover type in which they are located, (p. 135). A nest within a thicket is no safer from predators than is one located in a more open spot. (p. 136). Nests located within ten feet of an opening (including woods' roads) are no more likely to be broken up by predators than are those at greater distances, (p. 136). Grouse broods exhibit certain cover preferences as follows: The outstanding characteristics of brood cover are its diversity and its youth, (p. 139). Most used by broods are overgrown lands deficient in conifers, second-growth hardwoods, patches of popple, birch or alder and the more recently cut-over areas. I p. 1 40 ) . The presence of conifers contributes little to brood cover, (p. 140). Cover use changes, but not markedly, with the age of the brood. I ]j. 141 J. Broods prefer types in which the crown cover is sparse. I jj. 1 41 I. No type of cover is shunned by broods at any time of (ia\. Woodlands are preferred for night roosting, early in the morning and in late aflcriiooii. Overgrown lands and slashings are generally patronized during mid-morning. The former are also partic- ularly attractive during the mid-day and early afternoon periods. In late afternoon many birds seek out the alder runs and the young slashings, (p. 142). Broods tend to prefer the more open hardwoods when the weather is colder than normal and the types containing some conifers when it is warmer than usual, (p. 142). Wind but little affects brood cover choice. I p. 142). Woodland types are more used in rainy weather, overgrown lands and cut-over areas when it is clear, (p. 144). Ground conditions (wet or dry I seem not to affect materially brood cover choice, (p. 144). The more complex the undergrowth, the more attractive it is to broods. Small hardwoods, shrubs and berry bushes and a "combination of types"" are most frequented, (p. 146). Undergrowth density does not seriously affect brood cover choice. Sparse undergrowth is more frequented by the very young birds; medium and dense, as they grow older, (p. 147). Steep slopes are markedly less used by broods than are level lands or moderate inclines, (p. 148). No aspect of slope is shunned at any time of da) , although minor differences in use have been observed, (p. 149). Adult grouse behavior in relation to cover and related influences niav be sununarized as fol- lows:— Adults spend most of their time in woodland cover types, (p. 152). 108 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS In winter the birds frequent the more heavily wooded areas particularly if conifers are present, (p. 153). In spring the birds are still partial to woodland types. Excursions into cut-over patches and overgrown lands become frequent, (p. 153). In summer the birds revel among the overgrown lands, cut-over areas and in the spot- lumbered woods. At this season, types containing many conifers are seldom fre- quented, (p. 153). In early fall hedgerows and brushy patches, particularly if they border on conifers, draw birds like a magnet although all other types, save open land and mature hard- woods, may also be frequented. As the weather becomes colder, overgrown lands and cut-over areas are less used; woodland types containing conifers are more com- monly patronized, (p. 153). While few, if any, cover types are completely avoided even for a single nmnth, the degree of use varies considerably, (p. 156). Conifers represent the most used type for eight months out of the year but are avoided in summer, (p. 157). The number of grouse found in any one cover type at any particular time of day varies considerably with the seasons, (p. 159). Sunup is the best time to hunt the brushy pastures, woods' edges and cut-over areas in the fall. (p. 161). Weather does not usually exercise a strong effect on cover use. (p. 161). Birds tend to move into mixed second-growth hardwoods and conifers or to heavy ever- green cover when temperatures are colder than normal. Conversely on warmer than normal days birds move to the more open cover types deficient in conifers, (p. 161). Heavy conifers and alder patches (when the leaves are present) comprise the most sought out refuges on windy days. (p. 162). In snowy weather, birds seek coniferous cover, (p. 162). Rain bothers the birds but little. The more open woodlands arc tiien preferred, (p. 162). When snow covers the ground the birds are most apt to be found among the conifers while the mature hardwood type is preferred for snow-roosting, (p. 163). Grouse spend most of their time on the ground at all seasons of the year, though they frequently use trees for budding in the winter, (p. 163). Slope, degree or aspect, as such, seems to influence grouse cover choice but little. (p. 164). Most grouse flush within 50 feet of a person. Much-hunted birds generally flush wild as they do also on windy days. Females, gcnornljy. may be more closely approached before flushing than males, (p. 165). Flushed birds normally alight in the same type of cover from which they were originally started except when it is not extensive or is quite open. (p. 1661. Providing they do not remain in the same type of cover, wiien flushed, they tend to choose certain other types in which to alight, depending on the type of cover from which they were originally flushed, (p. 166). SUMMARY 109 Upon being flushed, birds normally fly only a short distance before alighting. Males do not usually fly a greater distance than females upon flushing, (p. 167). Two-thirds of all the birds flushed may be expected to alight in trees, (p. 168). The quality of a habitat is roughly proportional to the variety in the composition and arrange- ment of its component cover types, (p. 168). The poorer the covert, the greater the need for "edges." (p. 173). In highly productive coverts, such a relationship is less evident possibly because the varied composition within many types creates a diversification of cover, (p. 173). Solid blocks of coniferous reforestation are used but little by grouse except along the edges, (p. 174). Selective lumbering improves most grouse covers unless the conifers are thereby largely eliminated, (p. 176). Where forest cover is dense and extensive, the clear-cutting of small areas may profitably be used to provide summer and fall feeding grounds, (p. 178). To every hunter of grouse, fall brings a recurring puzzle — in what cover will one find the most birds? The answer is by no means easy, for grouse are hardy individualists, and expe- rience does not always point the way. But, with study, patience and careful correlation of such items as the type of cover, season, slope, time of day, wind and weather, certain pref- erence patterns can be recognized and catalogued. Many a bird-wise sportsman has con- sciously done this for his favorite hunting coverts, using such general terms as woods' edges, overgrown land, open land and evergreen thickets. In the more exact terminology of the investigator these cover groupings become vegetative cover types. It is his job to recognize and evaluate the multitude of natural climatic, physio- graphic and biotic factors and, through repeated observation, to uncover gradually the often complex part each plays in influencing the survival of the bird. The effect on grouse of each of these groups has been studied in some detail. The birds are tolerant of a wide variation of climatic factors — atmosphere, temperature and light. With a few exceptions, neither rain nor wind, snow nor drought, cold nor warmth, seem seriously to influence their survival. Physiographic features — slope aspect and degree — seem likewise of minor direct importance. Birds may be found in goodly numbers on steep slopes or flat lands irrespective of the compass direction the slope may face, providing the proper cover is present. Indirectly, of course, these factors usually play a commanding part with soil in determining the character of the third group, the biotic or plant and animal associations pres- ent. It is this group that largely decides whether grouse survive, become abundant or dis- appear. Let us carry this a step further. As between plants and animals, it is the former that are by all means more important in maintaining high grouse populations. Good cover means fewer losses from predators and probably from disease as well. In other words its "carry- ing capacity" is higher. One must, therefore, learn to recognize the important components 110 COVER CHARACTERISTICS AM) SHELTER REQUIREMENTS \\lii( h make up good grouse cover as the first step to intelligent management. it is. then, the purpose of this and the succeeding chapter to describe cover characteristics and to explore and define, in some detail, the shelter and food requirements of the grouse. While classified as woodland game, these birds find cover conditions to their liking only where a diversity of vegetation fulfills their food and shelter needs throughout each season of the year. The fact that the grouse is a sedentary species with a normal cruising radius of less than a mile and a minimum territory requirement of perhaps ten acres, suggests either very simple requirements or a fairly high degree of dependence on the occurrence of suitable habitat conditions within small areas. The latter surmise is more nearly the case for in large measure, it is the quality rather than the quantity of the cover that determines the number of birds a given woodland can support. Quality is seldom easy to recognize. One can determine the characteristics of grouse cover only by locating the various cover associations which support grouse and by finding just what part each plays in the life history of the bird. By stud) ing the use grouse make of the trees, shrubs, grasses and herbs, individually and as a conununitv. that comprise a cover type, one may gradually build a picture of food habits and shelter requirements. Man has, as yet, few ways of perceiving bird needs except by the roundaliout method of charting actions and. from ihem. attempting to recognize the compelling reason. The inter- pretatit)n is, of necessity, slow, for we have only our own experience as a guide and imagina- tion is required to put ourselves in the place of a grouse in attempting to explain its reactions. Because the probability of error in such observations is so great, a truly staggering number of these must be recorded covering each individual jilace and activity*. The number must be large enough to rule out the chance that, when anahzed. the result secured may represent but an incidental reaction to a situation rather than a characteristic habit of the species. Ruffed grouse range extends from Georgia to Hudson Bay and Alaska, from the Atlantic to the Pacific, from Long Island's low-lying plains to the stunted forests atop the higher Adir- ondacks. Throughout all of it the broad factors that govern grouse productivity are the same. But the specific combinations of conditions prevailing in different localities vary widely. This is particularly true of the plant and animal associations. Thus a study of the relationships of the species in New York can reveal its shelter needs as a whole but evalua- tion of the part each cover ty|)e plays in fulfilling them can be attempted onl\ fur liie situa- tions actually observed. CHARACTERISTICS OF PRODUCTIVE GROUSE COVER For practical purposes, one may divide grouse cover into natural groups such as open land, overgrown land, woodlands and slashings or cutover areas. Most of these, in turn, may be further sub(ii\ided into subgroups called cover ty|)es. Woodlands, for instance, may be composed of blocks of second-growth hardwoods. mMtini' liarduddils. cunifcrs or a mix- ture of hardwoods and conifers'^. Each large plant community and cover type carries within it ;i special set of conditions. Thdugli grouse are adaptable birds, they have by necessity become more or less specialized for li\irig within a fairh broad range of cover conditions. The extent to which they can find the basic life needs met within a given cover type is a measure of that type's importance and use- • Ttip ronrliisionii rrachril in this chapter mulled from a statiitiral analvtis of comprrlicnsivc data sliceta coverini! 19.619 i;rou»c fliDillra. 1,515 liruod rontarlH and llip rxaniination of 1,270 neat lorntinna. These records were nathered. over a period of 13 years, from all three iliverse types of grouse habitats in New York Stale, A See rover type descriptions, p. 120. CHARACTERISTICS OF PRODUCTIVE GROUSE COVER 111 fulness to the bird. What, then, are the characteristics of productive grouse cover? In General Food and shelter are the two prime requisites of life. The largely wooded habitat in which grouse live, must provide these conveniently and in abundance throughout the entire life cycle from egg to adult if the birds are to prosper. Each type of grouse cover normally produces some of both, but in widely varying amounts. Few, if any, cover types contain within their borders amounts adequate to supply grouse needs for a month or a week or. with some types, even for a day. Furthermore these needs vary with the kind of day, the season and the particular activities that are being carried on. For instance, in the winter, cover furnishing favorable shelter is a prerequisite, for the danger from predators and unfavorable weather is then at its height. Food, on the other hand, is not a critical item for much wooded cover is sufficiently varied as regards composition so as to provide the necessary buds and fruits. In sjjring the prime need is for types pro- viding adequate breeding and nesting cover. In summer, feeding grounds for the broods are important. Fall brings on the harvest moon for man and bird, with covers that support fat- tening foods receiving the lion's share of attention. These seem to be the outstanding seasonal requirements, though of course there are others of lesser importance. To meet these, good grouse cover must possess certain characteristics. Covert Size and Shape The smallest unit of fairly isolated habitat in which grouse liave nested, raised broods and maintained themselves the year around, to be surveyed by the Investigation, was about 10 acres in extent. Though completely surrounded by large open fields, birds occasionally trav- eled between it and other woodlands in the vicinity, thus making it impossible to state with certainty that, as a unit, it could have been self-maintaining. The same may be said of two other coverts, similarly situated, containing slightly less than 25 acres each. Providing the |)roper cover tyjjcs are present and productively interspersed, there seems to be no upper limit to the size of a good grouse covert. The shape of a covert, though often of interest to the hunter, seems not to be too important to the grouse. Long, narrow strips seldom provide for the proj)er interspersion of cover types. Rectangular-shajjed coverts provide more "outside edges", in which food is often abundant, than do areas that are more nearly square in outline. This is especially true where the coverts are disconnected and a])t to In' small as in many farming communities. Type Size and Shcpe The reverse of the situation just described is often true as regards the size of the various types that make up a covert. Mininnim dimensions are usually unimportant. A few clumps of a dozen conifers each may provide adequate winter shelter for several grouse. The cut- ting of a few "wolf" trees, with wide-spreading, dominant crowns, in a woodlot may let in light sufficient to encourage small patches of briers furnishing summer food for a brood. The important thing is that the type be large enough to create an "edge" and to allow for the development of the vegetation characteristic of that type within its borders. But maximum size is another story. The amount of "edge" is inversely proportional to type size. Furthermore large blocks of a single type seldom encourage maximum popula- 112 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS tions, since use, too, is often inversely proportional to the distance from the type edge. For instance, a study of the use made of blocks of coniferous reforestation* indicated that birds seldom penetrated much beyond 200 feet inside the block. Likewise, large, overgrown fields are seldom patronized much by grouse except where an edge adjoins wooded cover. One cannot, however, carry this idea to extremes. Each type fulfills a function in providing certain necessities. In general those that provide food (overgrown lands and cut-over areas) may well be small and scattered. Those furnishing shelter at critical periods (particularly in winter and during the spring breeding season) meet grouse requirements better if they are somewhat more extensive. GROUSE ARE SELDOM ABUNDANT WHERE THE FORESTS ARE EXTENSIVE AND UNBROKEN Much of the foregoing discussion is also equally applicable in considering the shape of a type. In general, the larger the "edge" the more productive the type, for the opportunity for grouse to make use of it is thereby increased. Irregularly shaped pieces are both desirable and the rule, though extremes may result in lowered use'''. Type Composition The vegetative makeup of a type is determined by climate, soil, slope, accidents such as fire, by the life within it and by man's activities. Grouse cover needs are best met by vegetation characteristic of the intermediate stages in plant succession from open land to climax forest. * Described on p. 174. A See Chapt, XIV. "Deeisniog Croute HsbiUti" for tuggested type pattern. CHARACTERISTICS OF PRODUCTIVE GROUSE COVER 113 Nature, often aided by axe and plow, produces the type; man distinguishes and classifies it according to its composition to meet his needs*. So also does the grouse. Though the makeup of a type may vary considerably, its essential vegetative characteristics are sufficiently constant so as to meet more or less completely certain requirements of the bird. Just how completely depends upon composition. A type reaches its best development for grouse when a large variety of species distinctive of it and useful to the birds are present. An uneven distribution of species and of plant density within the type is usually a sign that it will adequately fill its niche by providing certain neces- sities of life for the grouse. Likewise the more luxurious the growth (though not the density) of the various species, the better the net result is likely to be. A more detailed discussion of type composition will be found later in this chapter for com- position is one hallmark of quality in grouse cover. Interspersion of Types The same principles that make plant variety and interspersion so desirable within a type, apply with equal force as regards the arrangement of types within a covert. Two of these, furnishing essentially similar services, such as fall feeding grounds, adjoining each other, are less likely to be used than as though they were separated by types furnishing acceptable shel- ter and resting opportunities. On the other hand, should birds have to travel far from food to shelter over open land, for example, they would be more subject to predation than as though the types were adjacent. There is, then, a rather definite pattern of type arrangement which must exist if a covert is to be productive of grouse. The types that provide for the critical necessities, winter shelter, spring breeding grounds and summer and fall feeding grounds must lie within the territory'^ of, and be available to each grouse without necessitating the taking of undue risks or effort to reach them. The greater the distance any one is from all the others, the larger the territory occupied by an individual or a group of birds must be and the smaller the number of grouse the covert will support. The territory requirement of an individual grouse varies considerably. Psychologically a bird may be satisfied to maintain ten to fifteen acres as its especial domain, though at various seasons it may be shared with other grouse. But the cover quality and interspersion is often so poor that its food and shelter needs cannot be met without including 50 to 75 acres in its estate. A major objective in cover management, then, is to encourage the development of a type pattern in which these cover requirements are met within a relatively small area. No matter how productive an individual type may be. grouse will not use it to the fullest extent unless it is properly located with respect to other types. The order of arrangement seemingly matters but little; it is the fact that all necessary types are to be found within the normal cruising radius or territory, that is important. By Regions in New York State On the basis of grouse habitats three major regions may be recognized in New York. In figure 13 the portions of the State most typically representative of these are shown. While * See description of types recognized, p. 120. A See diecussion of grouse territories, p. 257. 114 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS the fuiiHariiental character of each is distinct, there are wide l)elts around and between them where they merge with and o\erIap one another. Also included in many parts of this interme- diate zone, particularly along the Ontario-St. Lawrence plain and in the Mohawk-Hudson Val- ley, are areas of intensively farmed agricultural land where grouse coverts are few and of low quality. REGION Adirondack ffi5.:S>!i Cats KILL 1. Connecticut Hill 2. Pharsalia 3. Adirondack {ijijjijjjjji Rest of State 4. Catskill FUaiRK 13. PARTS OF NEW YORK STATE MOST TYPICAL OF TIIK TllUFK MA.IOR (iROl'SK HABITAT REGIONS AND THE LOCATIONS OF THE INVESTIGATIONS STUDY AREAS IN EACH The basic factors controlling grouse abundance, such as shelter, food, predation and disease have been studied in some detail in <'a(h region*. The conclusions thus reached are there directly applicable. Nevertheless most of ihem are probably valid, in the main, for other sections of the Northeast where similar grouse habitats prevail. A brief description of the conditions in each mav, therefore, be of more than loial interest. The Adiroiiihicli Rctiioii The granite rocks and sands that make up the Adirondacks are among the oldest to be found on the North American continent. The soil is shallow, sandy and poor. The region is * T)ir litralionik nf ihc principal nludy arraa uaril tiy (iir Invraliliatinii arc sliown in ftKurt* l:t. With rraprrl to nciila, rcconla from fl«cwlirrp in tlic Stair liiivr lii-rn rtaniiifii-il an f(»lli>WH : nil cuunlir» rnvrn-il in whole or in |>art liv llir typical portion of a re- cion linvr lirrn inrluilf<] iiiidrr it. rxirpt thtil thi unties of Oneiila. fferkinicr nniitli of the Mohtiuk River. an<] Snratnca have been ron^iilereil part of tile Heat of Stale region ; all other counties have been jilaeed in the latter region also. CHARACTERISTICS OF PRODUCTIVE GROUSE COVER 115 characterized by large tracts of forest, unbroken, except where rock outcroppings or wind- falls provide occasional small openings. Interspersed among these areas one finds beaver meadows, lake shores, alder beds, cranberry bogs, burns and. about the settlements, clearings. Mountain slopes and swamps were originally clothed with spruce, balsam, pine or hemlock mixed with hardwoods, notably beech, birch and maple. Although some virgin timber still exists, chiefly on State land, most of the region has been cut over. The larger conifers and some hardwoods have been taken for timber, the smaller ones floated to the paper mills for pulp. The few roads traversing the area have made lumbering costly except on an extensive scale, thousands of acres occasionally being involved in a single operation. On the higher ground the succeeding second-growth is predominantly hardwood with spruce or pine while in the lowlands and swamps, spruce, balsam and larch are often abundant. Here the white- tailed deer have repeatedly browsed off the smaller hardwoods and the balsam. Largely as an aftermath of logging, a number of severe burns have occurred, particularly in 190.3 and 1908, creating barrens into which such pioneer species as pin cherry and aspen are gradually seeding. In terms of providing for the c<)\er requirements of grouse, most of the Adirondack forest leaves considerable to be desired. Both second-growth and cut-over land are apt to occur in large blocks, thus affording little of the interspersion of types commoidy found in the other regions of the State. Conifers affording winter shelter are abundant, but the herbs, berries and fruits furnishing good summer and fall food commonly occur in numbers only in the alder runs, along the edges of the burns, in the recently cut-over lands and in the small, often grassy, openings resulting from windfalls or from one or another of man's activities. In such places grouse tend to concentrate; elsewhere they are usually quite scattered. About the periphery of the region, however, the soils are more fertile and, as the forest cover becomes more and more interspersed with farm land, one finds habitats of much better quality. The Catskill Region Although similar in many respects to the Adirondacks. the Catskills have a forest cover more varied and less continuous. Mostly a region of sedimentary rocks, its forests are of birch, beech, maple, jiine and hemlock, with spruce at higher elevations, particularly on the northern slopes. In type thev are more closely associated with the Alleghenies to the south and the lower New England mountains to the east. Pastures thrust long fingers up from the farms scattered along the valleys. The region has been extensively cut over, mostly in rela- tively small blocks, the conifers for luniluM- and the hardwoods for cordwood and for saw logs. In the western part, where acid wood distillation was once a thriving industry, the slopes are largely grown up to second-growth hardwoods interspersed with hendock. Much of the woodland is grazed though seldom with sufficient intensity to encourage much grass. There are many overgrown pastures. The soil is more fertile than in the Adirondacks. thus producing a greater variety of species and density of cover. Where the crown canopy of the woods is not too thick, undergrowth and ferns, including mountain laurel and rhododendron in the southern half, occur over large areas. Openings and cut-over lands commonly support a profusion of herbs and berries. They also seed in fairlv rapidly to hardwoods and. occasionally, to hemlock and pine as well. Abandoned farms are not uncommon. As might be anticipated under such conditions, woodland, cut-over areas and brushy pas- 116 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS tures often combine to furnish much high quality grouse habitat, although, along the ridges and at the higher elevations, the continuous forest cover seems to meet less satisfactorily the varied needs of the birds. Rest of State The regions just discussed contained much land too steep, too stony or too difficult of access to attract many settlers but, in their broad periphery and across the south-central and south- western part of the State (locally known as the .Southern Tier), conditions were more favor- able. On the hilltops farmers moved in, cleared or sold off much of the pine and hemlock, ►■ THE ADIRONDACK REGION CONSIST.S OF MOUNTAINS AND WIDE LOWLANDS, FORESTED CHIEFLY WITH SPRITE. BALSAM. PINE. BIRCH AND MAPLE. IN THE VALLEYS ARE SCATTERED PONDS, SWAMPS AND CLEARINGS mined the shallow top soil of sidehill and upland and stripped the woods of everything worth selling and moved out a century later. In the deeper valleys and occasional fertile upland pocket, however, farming has prospered. Here woodlands have shrunk to woodlots. held as a source of lumber, posts and cordwood needed about the farm. Where these arc sufficiently large and not too heavily pastured, some grouse usually find acceptable, if limited, coverts. But in the poorer uplands, where the farmer has given up or is at best making scant prog- ress in combatting the shrubs and trees which seem, in a few short years, to take possession of his unworkcd fields or his jjoorly grazed pasture, grouse habitats, as productive as any in the Northeast, are commonly to be found. Here Allegheny and Northern hardwood types meet. Oak and maple, beech and birch, pine and hemlock vie with each other in endless pro- fusion. The partly abandoned farmlands and pastures are constantly being overrun by fall THE CATSKILL REGION IS A LAND OK NARROW. FARMKl) VALLEYS WITH PASTL RKS E\rtM)lN(, IP THE SLOPES TO MEET EXTENSIVE FORESTS COMPOSED MAINLY OF BEECH, BIRCH, MAPLE, HEM- LOCK AND PINE THE REST OF STATE REGION IS CHARACTERIZED BY FERTILE VALLEYS AND POORLY-FARMED UP- LANDS. HERE THE MOSAIC OF ABANDONED FIELDS. OLD PASTURES AND WOODLOTS FORMS MANY A PRODUCTIVE COVERT 118 COI ER CHAR ACT ERISTICS A\D SHELTER REQUREME.XTS grouse food species such as pin cherrv. aspen, thornapple. wild apple, sumac, dogwoods and \iburnums. These commonly occur adjacent to woodlands in which hardwoods and coni- fers furnish winter shelter and spring breeding grounds. Woodcutting operations still make a\ailable a profusion of herbs, berries and insects for the summer use of the broods. The extent of any one covert is seldom large, thus providing a characteristicallv high inlersper- sion of cover types and a wealth of desirable grouse habitats. This is but a \ignette of New York State grouse coverts. It is the setting in which the Investigation carried on its work. Detailed descriptions of the physiographic features and plant associations of each region have been purposely omitted since a chapter would be necessary to describe each in detail. THE ROLE OF SUCCESSION In outlining the makeup of grouse coverts in each of these regions emphasis was intention- ally placed on the forest uses, past and present. The extent to which such crops as forage or wood products are harvested may affect the productivity of a woodland for grouse quite as much as does its natural composition. Most of these practices tend to arrest or throw back its normal development to an earlier stage. This may exert a strong influence on the grouse, for each stage varies considerably in its ability to produce the required food and shelter. Likewise grouse need change with the seasons as well as with the age and the sex of the birds. The partridge has justly been called a transition type species in that it finds its year-round requirements largely met neither in the first stages associated with grasslands nor in the last or climax tvpes of dense unbroken forest. The productive coverts are those in between. The jirogression by which one stage takes over and grows into the next higher association of plants has been termed succession. But. since in grouse habitats considerable time is in- volved in making the change, many hunters fail to recognize in this a likely reason why there may not be as many birds in their favorite coverts as they once remember. Grouse cover is constantly changing. Insects and disease, fire, the axe. cow and plow alter the normal succession. \^Tiere the climax type is forest, meadows, if left alone, seed in sooner or later to rank herbs and brush. The lime and kind depend on many things. Among these are site, seed supply, weather and the activities of birds and mammals, particularly mice. Quick-growing species that seldom do well under shade usually thrive here, furnishing a shelter under which the more permanent species find growth conditions favorable. Manv of the larger fruiting trees and shrubs, such as sumach, cherry, apple, thornapple and grape, require considerable sun if thev are to bear abtnidantly and. accordingly, reach their lic«t de- velopment on such overgrown lands. On the degree to wliii h ihis development has progressed depends the use grouse make of such overgrown lands. As long as the vegetation remains open and luxuriant in character. an abundance of insects, seeds, fruits and desirable leafy material characteristic of such cover attract grouse and adults throughout both the summer and fall. Once the trees take over, shut- ting out the sunlight, most of these plants gradually disappear. Its imjiortance to grouse changes accordingly. Left alone, overgrown fields e\entuall\ become second-growth woodlands with a more or less dense undergrowth or a ground cover of herbs, sedges and grasses, depending on soil fertility, moisture and the existing plant association. Gradually, the quick-growing, sun-loving species of the previous stages are shaded out b\ the taller oaks, beeches, birches, maples and A-I^. IM X u D3 U -] o X a: a. C o X a o o u .f - ,^tiiJJ*i a: yWK c z < o O 2 c H K >±^ > o U c z f- Q O a U > U! Q -;; ,< U m u rn >- a -? o ^ fi] III en S-£ ^^'v o z o z <; J H Z -s H Z o u -*>.iLtr- 1 :i c Z O %1 M O 120 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS conifers that replace them. WTiere the crown cover is sufliciently dense to maintain fairly open conditions beneath, such woods are often chosen as spring nesting grounds; where they are more open and the herbs and ferns thicker, they furnish considerable summer feed. With the addition of clumps of shade-loving conifers, such as hemlock, spruce or of the more light-demanding pines and balsams (the latter usually started while the land was not yet heavily wooded), the require- ments of the grouse for shelter, especially in winter, may also be met. As the forest matures and closes in. barring action by Nature or man to create openings, most summer and fall feed gradually disappears. The birds are then forced to seek a more congenial habitat along the edges. Let a wind storm create openings by uprooting the less se- curely anchored trees, or a lumbering operation open up the cover, and the succession is set back accordingly. In the openings thus created herbs, briers and berries quickly spring up thus again providing abundant feed for old and young alike. Then the birds move in again. Thus goes on the ]iageant of succession. By it, the grouse are pushed hither and yon, ever seeking the places where winter shelter, spring breeding grounds and summer and fall feed lie close together. On these they must depend to a greater extent than is generally realized, for the essentials of their existence. In these they find varying degrees of protection from their several enemies whose success depends largely on finding the birds in the poorer cover. COVER TYPES RECOGNIZED Botanists and foresters divide the various stages of succession through which a woodland may pass into cover types. The hunter, perhaps unconsciously, long ago applied the same technique as an aid in finding his birds in the fall. The basis of recognition, in one case, is largely what prows there; in the other, the use the birds make of the plants. Thus a scat- tering of thornapples. cherries and dogwoods along a wood's edge becomes known as likely fall feeding grounds for the grouse. The emphasis is here clearly and rightly placed on use. But the intricacies of cover are such that no clear picture of use can be olitaiiied from chance observations alone. Neither is it practical to classifv cover into innumerable small subdivisions, for the number of records needed to determine to what extent each fulfills grouse cover requircrncnls is too great in the face of the varied habits of the bird. Based on experi- ence, the Investigation gradually came to recognize the following twelve major cover types, representing four major stages of succession. (Table 13). TABLE 1.3. EXPLANATION OK THE C0\ EM TYPES AND SYMBOI>S USED L\ THE RUFFED GROUSE INVESTIGATION. Stages of succession Typp of cover Type rode Open Lnnd A OvfrRrowii (.jincls R C D Wo<»ltiiri.|H Prodoininiiti-ly hiinlwnod.H (iU'M)% conifers). Llirndy second growth, mostly .■J'-I2'' in diameter. . . Mixed CM-~i}%) hardwcKids or conifers. Lnrpely second proutli. mostly If-l^' in diiimeter IVetlomiriiilely hiirdwiuMls (0-'M)% ronifers). I.nrtrely miitiire, ninstly over 12' in diumcler K EH F FH Sf>(»t-liindH're(|. \i)-7i()'/'g, of the crown eanopy removed liy si-lei-tive !<»>;«'"? G 11 I SlnahinRS J COVER REQUIREMENTS OF THE GROUSE 121 This type selection obviously is not orthodox by such standards as foresters have proposed. A botanist may shake his head doubtfully, though the choice may make better sense to plant ecologists. Perhaps only the grouse-wise will recognize the importance of emphasizing, as the essential difference, the character rather than the kinds or species of plants as the basis of type differentiation best suited to the purpose at hand. Any one of the conifers or the broad- leaved evergreens, such as mountain laurel, may represent, for a grouse, potential sheher from predators; a hundred different plants that grow in slashings may furnish it with food. In other words, the types here selected are largely representative of the different stages of forest succession or regression which function to fulfill some niche in the broad panoply of grouse cover needs. In following such a line of reasoning, it is easy to lean over backwards. One cannot simply ignore all species of trees or shubs in evaluating the whole, for some are more productive of food and shelter than are others. The grouse is, however, so adaptable that no one or two species are indispensable to its welfare. Rather it is the groups of plants more or less char- acteristic of each type, which, in whole or in part, serve to fulfill one or more of the bird's principal needs. The degree to which these are met is regulated by the composition of the group and the proportion of the whole which it occupies. This, in part, explains why grouse are likely to be found more frequently in certain parts of a type than in others. Other equally important reasons, notably cover arrangement, are mentioned later. The properties in each cover type that determine its value to the grouse may be spread throughout or localized here and there. Examples of the uniform situations which may occur are pure coniferous stands resulting from artificial plantings or an alder thicket so dense as to preclude the intrusion of other light-demanding species. Conversely, in brushy pastures, the best food conditions for grouse are usually to be found at some distance from the gate. Though the details may differ, counterparts for most of the situations here described are probably to be found throughout the range of the ruffed grouse, for though the species may differ, the pattern of woodland succession is much the same. Nevertheless, the usefulness of these particular type groupings in studying the shelter requisites of the bird or. in fact, of adapting many of the conclusions here drawn to other regions is dependent upon the extent to which conditions are similar to those pictured for New York and upon the success with which the influence of any differences can be evaluated. COVER REQUIREMENTS OF THE GROUSE Wherever grouse hunters gather, discussions are bound to arise. Where are the birds most likely to be found on warm October days or in a November snow squall? Is it worthwhile hunting steep slopes? How far will the birds wander from sheher in search of food? Thus the questions go. Occasionally the thought is expressed that, if the answers were definitely known, it might help an interested landowner or a state game department to manage some woodlands so as to produce more grouse. The only way to settle such matters is to turn to the birds themselves for the answer. It was with this thought in mind that the Investigation early started an intensive study of grouse cover requirements. Beginning in 1930 two primary study areas were established in the Rest of State region and one each in the Adirondacks and the Catskills*. Secondary units * The principal onrs wi-rr ih^ Conneclicul Hill. Pharsalia. Adirondack and Calskill survey areas. N a. Q Z < o ce o a: V 5 CL. H OS u b Z o a c o ce u Q z o a o o -'•■»-«*W* ViiSfm^"*^- ■••'At. y- X U [I. Z o c o c Q a o o a. e u u. z o o p. < u b: ■< a u OS u « 3 .J H O a. a. ►J u u o S u X a u z o u 0. Q 2 C b] K H u b 2 O 120 COl EH CHARACTERISTICS A.\U SHELTER REQLIREME\TS were set up as needed for special project study. On the |)rimar) areas, surveys were in process for from six to twelve months over a 13-year period. Over lO.UUU man days were spent gath- ering data on the conditions surrounding the 19,619 grouse flushes, 1,515 grouse broods and 1.270 nests, which are here analyzed. The nesting studies were extended to cover those lo- cated by cooperators in most of the counties of the State where the grouse are common. Not being certain as to just what details might be of value, the policy of recording every item, uhirh might conceivably be of interest, was adopted. In furtherance of this, each time a grouse was flushed it was found desirable to record data covering 36 sej)arate items on a specially devised form sheet*. Similar notes were made for nests, broods and dead birds en- countered. By 1937, the mass of data thus gathered had become sufficient (over 700,000 records) to warrant its tabulation'^ as a basis for analyzing the shelter and other requirements of the grouse in New \ ork State. At the beginning, the authors little realized the complications involved in securing records representative of true conditions and sufficiently numerous to remove, in large measure, the probabilitv of error. Though the best statistical advice possible was earh secured, the ap- |)li(alion of biometrics to such an ecological problem was still in its infancy. Full advantage has been taken, however, of the tremendous advance in the past decade in setting up and evaluating the tabular material from which most of the conclusions later described, have been drawn. The problem of securing records representative of the actual situation was attacked by at- tempting to secure a sampling of actual conditions sufficiently large to reduce the probability or error to a minimum. Technically, representativeness can only be accomplished, as Dr. Mottley^ aptlv put it, by "giving each and every grouse nest, brood and adult, in every type of grouse habitat, in each year and under each condition, an equal and independent chance of being observed." Obviously, this is not practical in a species like the grouse although it was ap])roached in studying the cover relationships of grouse broods and adults by following the birds on most of the study areas, throughout all four seasons year after year, with about the same degree of ititensity. In arriving at the conclusions here presented, wherever practi- cal, the corrections necessarv to account for such items as variations in the amount of each cover type available to the grouse, have been made. This is necessary before one can secure a true picture of grouse cover use. In studying grouse nest locations, it is assumed, on the basis of experience, tiiat cacii bird will choose the cover type and nest site most nearly fulfilling its requirements. Many instances have been observed where females have travehxl considerable distances, presmnably to .secure conditions favorable to the location of the nest. Except for this special situation, it is believed that grouse I broods and adults) to a considerable extent must make the best of conditions within their territories as they find them. Thus the results here presented cannot be consid- ered to be absolutely representative of what a grouse WDuld select if given an unlimited choice or if all habitats represented optimum conditions. The lar^c rminl)cr of records analyzed, however, is believed suHiciciit to present a picture of "average choice" undi'r New York conditions. Nesting Cover Though grouse are not particularK adept at hiding their rii'sl. their concealing coloration. * Sop riniirrn 73 In 76 in Aiiprnilix. A Mptlliiilit liarit arc (lescrilird in tile Apprnilix. p. 701. t Mottlfy, C. Mi-C. pcrsunul Ivlirr to the author*. COVER REQUIREMENTS OF THE GROUSE 127 coupled with the habit of "sitting close"' when approached, makes the finding of one an event to be remembered. Thus it is not surprising to find no extensive studies and but few opinions expressed as to what constitutes good nesting cover. Are thickets or open spots preferred? Do the birds avoid steep slopes as nest sites? Of some 39 per cent annually destroyed* are those located in any one cover type or other situation most likely to be broken up? Such problems can seldom be solved with finality and, at best, only for the regions studied. The answers here suggested are based on a study of 1,270 nest situations, yet it is certain that, with constant improvement in technique and with more intensive study, others will add much to the picture. They may even prove some of our interpretations wide of the mark. Therein lies the fascination of wildlife research. It is sometimes easy to dictate results by the wa\ in wliicli one carries on an investigation. Of the nests here reported on, almost half wire f<]und wliilc making the spring sur\e\s on the various study areas. On these all cover types are searched for nests with equal thorough- ness'^. Most of the remainder of the nests were reported by cooperators or other iiuli\iduals who happened to stumble on them during the day's work. "Building fence" and "going after the cows" were the most productive occupations in this respect. Let us then look at some of the outstanding preferences relating to nesting cover. Types of Cover Chosen The distribution of nests, according to the frequency with which each cover type is used, is presented in table 121 ^ By studving it. one finds some interesting correlations. * Si-P Cliai.ur \ 11. |.. -111. A Fach area was comidclrly ruvcrrd onci' each wrck I'V thr Mir*f> iirw, Srr MfllnuU ami Trrliniijiir!. (p. 699.) t This and other tablca sunimarizine shelter relationship data haw been [ilareil in the .Apt'endil tteginning on p. 783. NEARLY HALF OK THK GROUSE NESTS IN NEW YORK STATE ARE TO BE FOUND L\ SECOND-GROWTH HARDWOODS (TYPE El I2ii COVER CHARACTERISTICS AMJ SHELTER REQUIREMENTS While some nests have been found in each type, nieadowlands (type A) supporting occa- sional trees or shrubs, and coniferous cover (type Hj are least utilized for this purpose. Woodland types are strongly favored as nesting cover, 899 out of 1,270 nests being found therein. Of all the woodland types, nests are most likely to be found in second-growth woodlands (types E and EH), for 565 (48.4 per cent) were located there. They are least likely to be placed among conifers in which but 52 (4.4 per cent) were discovered. Only about one-third as many birds are likely to place their nest in mature woodlands as in second-growth cover. Except for mature woodlands, which are apt to be fairly clear of nmch undergrowth or ground cover, all of the most likely cover types chosen are characterized by providing con- siderable summer food for broods. This is especially noticeable with respect to spot-lumbered areas (type G), which, although small in extent b> comparison with other types, attracted 105 birds (9.0 per cent). There is a tendency for the nests to be located in types deficient in conifers. Thus three times as many birds nested in overgrown fields deficient in evergreens (type C) as located their nests in overgrown fields where a substantial number of conifers (type D) were present. A somewhat lower ratio is evidenced between second-growth hardwoods ( type E), as con- trasted with a mixture of hardwoods and conifers (type EH). As the stand approaches ma- turity, this difference tends to become less and less. One finds second-growth hardwoods predominantly used as spring nesting grounds. The primary reason for this is still to be determined. The presence of considerable summer food and the more open character of undergrowth and ground cover in this type, in contrast with that existing in most overgrown lands and slashings may furnish a clue. Density of Undergrowth at Nest Site It is interesting to note from table 122* thai most birds apparently are reluctant to place their nests where the undergrowth is dense. Thus we find 40.4 ])er cent of the nests situated where the undergrowth is sparse and 46.4 per cent where it is medium, whereas but 13.0 per cent of the nests are located in dense clumps of undergrowth. The table suggests an interesting but not always consistent correlation between the density of undergrowth surrounding the nest site and the characteristics of the woodland in which the nest is found. In types where the undergrowth is normally rather dense, such as overgrown land, second-growth woodland, cut-over areas and spot-lumbered woodlands, a large prepon- derance of nests occupies the more open spots. Conversely, where the crown cover is hea\ \ and llic undergrowth normally sparse (such as in mature hardwood or coniferous wood- lands I the occasional more dense undergrowth clumps arc rather often chosen, though the inajority of nests are still located in comparatively open situations. There is a certain, though, when recognized, not serious error, wliich is clearly brought out by the above comparisons. Whereas densitv in all tvpes should ha\e been cdtniiared to a single rigid standard, in realitv there was a tendency to depart somewhat from this by considering tlic normal undergrowth for any type as medium. This may help to explain why. in cover tvpes characterized bv heavy undergrowth, grouse were more a])t to nest in the more open situations with the reverse being true in the types comparatively bare of undergrowth. * Sec Apprndix. p. 784, MANY A NEST IS PLACED AGAINST A STUMP OardiritT Bumji THE SITE MOST COMMONLY CHOSEN IS AT THE BASE OF A TREE Robert jr. Darrotc A VERY UNUSUAL LOCATION IS IN THE HOLLOW OF A STUMP l.iO COIKR CHARACTERISTICS AM) SHELTER Ri:(JUREME\TS It all serves to strengthen the idea that grouse prefer a rather open cover in which to nest. The word preference is used advisedly for here there seems to be a clear example of the exercise of choice by the birds. Certainly the undergrowth densities selected were not those most coniinoiih found either in overgrown lands, in slashings or in mature woodlands. Sites Preferred as I^est Locations The o])inion generally held that grouse are more likely to nest at the base of a tree or stump is borne out by an inspection of table 123*. Of the two sites, the base of a tree was chosen b\ 599 (51.7 per cent), whereas the next most attractive location, the base of a stump, was chosen by but 175 115.1 per cent). Nearly as many, 158 (13.6 per cent), located their nests under a pile or tangle of dead brush. The only other site coninionlv utilized was beside or tinder a log, where 120 nests ( 10.4 per cent) were found. Neither rock ledges nor clumps of bushes were |>arti(ularl\ poiJular although both were utilized. Interestingly enough, seven nests were found in c;Mties in stumps or in semi-roofed- over depressions on steep banks. Three were even located in oi)en fields with only such small shrubs as low-bush blueberry for jjrotection. A study of the sites chosen again bears out the conclusion that dense cover around the nest is seldom desired. Because of the concealing coloration and habits of the bird ])reviously re- ferred to, attention was called to most nests only when the female flushed. No difference was observed in the readiness with which a bird would flush from nests found in dense cover as compared with nests located in the open. Nest Location in Relation to Degree of Slope Since an entire grouse covert may. upon occasion, lie largely on but one slope, there does not always exist an opporlunitv for birds to choose a particular degree of slope. The same ajiplies with regard to the direction it faces, technically called aspect. For this reason, in analyzing table 124^^ any comjjarison of the number nesting on flat, gentle or steep slopes is subj<'(t to reservations. It was not possible to correct the figures in accordance with the proportion of each shipe available. Therefore it is best merely to indicate ihal. if occasion demands, birds will nest not only on flat or gentle slopes, but on steep banks as well. Since 281 nests 124.9 per cent) were found on flat land as compared with 149 (13.2 per cent) on steep slopes, one might conclude lliat it is the availabilit\ of the slope uitliin llie habitat, rather than its steepness, which largely inlluences the number of nests found in each category. Where gentle slopes are available, they are much utilized as is evidenced by the finding of 702 nests thereon. The predominant slope on most of the areas studied, however, lies in this classification. !\'est Location in Relation to Isjicct of Slope The same objection raised with the |ire\ious table is equally valid with table 125+ de- scribing the location of grouse nests as influenced 1)\ t\ pe of co\er in relation lo direction a slope may face. In addition, the difference in the rnunlici of nests foinul on a noitli or an east slope coni|)ared with a south or a west slope is so small as to dictate llie conclusion that no slope is shunned. Whatever advantage there may be is apparentK in faxor of west slopes which were utilized somewhat more frequently for nesting iIkui was an\ other aspect. None of the four slopes were utilized as freiiucnlK for nesting as was ilal land. * Sec A(»|n'inlix. (t. THS. A Sec AppciKlix, [I. 78(>. t See Appendix, p. 787. A NEST IS OCCASIONALLY SITL \TK1) BY \ PILF. OF LOGS Roherl il . Dirron A FEW BIRDS PLACE THEIR NESTS UNDER LOGS 132 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS It is interesting to note that no one cover t)'pe seems to be substantially more attractive than another when associated with any given aspect. No justification can here be found for the thought occasionally expressed that warm st)uthern slopes are most likely to attract nest- ing birds. Conversely, the more mature woodland types, offering greater shelter, are not pre- ferred on sunny slopes, nor are the more open types on northern exposures. In this connec- tion, one sometimes forgets that grouse establish their nests before the leaves are out and that, except where conifers are present, there might, therefore, apparently be no great difference between types in their sheltering effect on the nest. Undoubtedly some preferences do exist. Perhaps 1,270 nests are too few to bring these out. particularly when many grouse find no more than one or two aspects of slope available within their normal territory. Perhaps, also, were it possible to conduct a studv of grouse nest locations in a region where each aspect supported a different climax type, a valid difference in use might be obtained based not so much on exposure as on the differences in cover type occupying each aspect. For instance there is a tendency in southern New York for southern slopes to support a larger proportion of the oak and hickory type in contrast with beech, birch, maple and hemlock on the colder northern aspects. On the other hand, if all aspects contained the same climax types, then it might be possible to determine the effect of exposure on cover choice. With both factors known, the data could be more accurately correlated to find the relation of each to nest location. The Influence of Conifers on Nest Location It has already been noted that only a few grouse nests were found in predominantly coni- ferous cover types. Where hardwood types prevail, in the region studied, it is normal to find a considerable number of individual or of small clumps of evergreens scattered through the woodlands. Analyzing the records to determine whether these trees are particularly sought out as nest sites (table 126)* one finds but 181 out of 1,088 nests, located at the base of conifers. An additional 197 were found from one to ten feet away from a conifer, although often at the base of some other tree. The average distance of a nest from an evergreen was 136 feet. The fact that 228 nests were located more than SOO feet from anv evergreen, again indicates that conifers do not repre- sent a necessary element of nesting cover. Effect of Openings on Nest Locations Openings in the cover play a much larger part in determining the location of grouse nests than is generally realized (table 127)"^. One may spend many an interesting hour in the woods looking for nests but seldom finding them. The fact that 18.6 per cent of those located were in an opening while 56.0 per cent were within 100 feet of an opening of some description may help to explain one's lack of success while coursing the deeper woods. The word open- ing, however, must be interpreted broadlv. It includes not onlv meadows and fields border- ing on woodlands but also overgrown lands (types B. C and D) and recently cut-over areas which are still largely in the herb and berry stage (type I). Old woods' roads and trails make an often unnoticed break in the crown cover but provide a better chance for a somewhat greater variety of herbs and sometimes berries to thrive than is found in the surrounding cover. A bordering second-growth of hardwoods (type E) represents favorite nesting cover. • Spe AppendiK, p. 789. A See Appendix, p. 790. ■■v«-<«.-^j,,^ *„, -sV ^^o^ A LOOSK TANGLE OF BRUSH OFFKRS A LIKF.LV SPOT FOR A NEST >x ■■•^ BUT FEW NESTS ARE PLACED WHERE THE IMMEDIATE COVER IS DENSE 134 COVER CHARACTERISTICS A\D SHELTER REQL'IREMEXTS The general principle may be stated thus: — other factors beinp; equal, the desirahilitv of a nesting site varies inversely with its distance from an ojiening. This rule usually holds good irrespective of the type of cover in which the nest is located. In all types, the decrease in the number of nests as the distance from an opening increases seems to occur at approximately the same rate. Thus for 929 nests on which this point was specifically observed and which were not situated in an opening 197 (21.2 per cent) were found within ten feet of open cover and 261 (28.1 per cent) in the next 40 feet. Out of the whole group, in only 143 in- stances (15.4 per cent I . was a nest more than 200 feel frcini an opening. The probability that many nests will actually be located in some of these openings has already been mentioned. It is astonishing to see how close some birds will j)lace their nest to well-traveled roads. Two instances, in particular, come to mind. One was on the southern (lank of the Catskills, where a bird placed her nest on the overgrown hank of a steep cut through which a concrete highway passed. In the other case, the nest was found under a small shrub a scant six feet from a macadam highway in the northern Adirondacks. The application of this principle of proximity to an opening in designing productive grouse habitat is covered in some detail in the chapters on Management. The Effect oj Slashings on \ I'sl Location Of all the oi)enings, no one type normally contains as much summer food for grouse broods as do recently clear-cut areas. For this reason, special note was made of the various dis- tances from a slashing at which nests were located in the different types of cover (table 128)*. The data indicate clearly that nests are more likeK to lie Iik alcd within tlie first ten-foot strip of nesting cover surrounding a slashing than in llie scciukI. the tliird nr am iimre dis- tant strip. In comparison uilh o])enings in general, however, (table 127)'^, the inniicdiate vicinitv of a slashing does not seem to be substatiliall\ more attractive. One finds n<> greater tendency for the nests located near these units to be (loser tlvrctii than were nests located near Dtlui t\pes of o|)enings. It nnist nut Ijc lorgotten that voiuig grouse are precocial bird.--, tlial is. the\ take care of themselves to a large extent IKmi llie lime they are hatched. This includes linding practi- cally all of their own food ariil. if it a|)pears desirable, traveling relatively long distances soon after hatching. This was evidenced by one brood, banded at the nest and contacted 24 hours later more than a half mile froTn that point. Other broods, not disturbed, have moved lesser distances or been contacted, for several days after hatching, in the innnediate vicinity of the nest site. Some grouse locate their nests close to another opening even though a slashing ma\ be in liie iiinuediati' \irinit\. To sum up, a slashing seems to be a desirable luit li\ rm niean> necessary adjimet to good nesting cover. This ty|)e really comes into its ow n as a piddurer of sunnner and. to a lesser extent, fall foods for young and adult alike. In/Iiicncc oj Other T\/ic\ oj Ojicniniis on \cst Location 0|»eMings are like magiii-ts of dilferent strength uliieh ser\e to draw nesting grouse 0. COVER REQUIREMENTS OF THE GROUSE 135 or into them. To this conception, table 129* lends considerable weight. When this table was analyzed, it was a distinct surprise to find more than a third (41.6 per cent) of all nests, that were not actually in an opening, located within 100 feet of a road or trail. This figure seems high unless one remembers that two of the three regions of the State are, to a considerable degree, farmed. Most woodlands adjacent to farms have several more or less used woods' roads or trails within their borders. The 100 per cent coverage given each study area by the survey crew, however, precluded any chance that the cover along woods' roads might receive more than its share of attention in looking for nests. Of the other types of openings, grouse are most likely to place their nests not far from open land. In total numbers of nests, this situation outranks both slashings and overgrown lands. Young slashings seem to furnish more attractive conditions alongside which to locate nests than do overgrown lands. As previously mentioned, the attraction of a road, trail or young slashing apparently di- minishes rapidly as the distance therefrom increases. This tendency is much less marked with open lands. The nests are almost as likely to be placed at any distance up to 100 feet from an open field as within the first ten feet. Effect of Crown Cover on Nest Survival Having seen the effect cover may exert on nest location, let us examine the difference it may make on the success of nests |il;i will be seen in table 130^ 43.5 per cent^ of all the nests located were broken up. Any relationshi|) between cover and the like- lihood of nest destruction may, therefore, prove im|)ortant. As indicated in the table, cover seems to bear little relation to the fate of the nest except in a few cases. Nests placed in stands largely composed of conifers (type H) are most likely to be broken up. Conversely, those located in selectively logged woodlands (G), with its scattering of lush herbs and undergrowth where, here and there, a tree has been cut, are least likely to be found by predators. It is probable that this difference holds in all regions of the State though this cannot be inferred from the table, for the numbers, thus divided, are too few to show significant dif- ferences except for the Rest of State region. Here the trends follow those already described although there is possibly a greater likelihood of nests in a mixture of mature hardwoods and conifers also being left alone. Effect of Undergrowth Density on Nest Survival As with crown cover, it is remarkable how little real difference the density of the under- growth seems to make on the fate of the nest (table 131)^. Of 268 nests found in sparse un- dergrovrth 60.4 per cent hatched; in medium density, of 300 nests, 59.0 per cent hatched, whereas of 95 nests found in dense undergrowth 55.8 per cent were successful. When the fate of grouse nests is correlated with undergrowth density in the various types of crown cover, no significant tendencies are likewise to be found. There is a slight indication • See Appendix, p. 792. A See Appendix, p. 794. t This figure differs slightlv from ihe averages indicated in Chapters VII and XII since it is based on records prior to 1937 while the latter include the period through 1942. and also because shelter data were included for certain broken-up nests found while collecting eggs for propagation. X See Appendix, p. 796. 136 CUl EK CHARACTERISTICS AND SHELTER REQUIREMENTS that nests are safer in sparse undergrowth in second-prowth hardwoods (type E). The same appears likely in the open undergrowth beneath mature hardwoods and conifers (FH). But the inference is not strong and is further weakened when the companion types (EH and F), upon examination, are found to show this tcn(lenc\ in reverse. Thicket versus Open Cover as NesI Locations As has been seen, grouse prefer to locate their nests where the undergrowth is neither par- ticularly dense nor open. Referred to above, however, is the character of the undergrowth in the general vicinity of the nest rather than that actuall) at the nest site. Table 132* was accordingly prepared to answer the occasionally raised question, "Is a grouse nest, without benefit of immediately surrounding cover vegetation, more likely to be broken up than one located in a thicket?" The answer seems to be that it is not. Of 630 nests with but little sheltering cover, 362 (57.5 per cent) hatched, whereas of 208 thicket nests, 110 (52.9 per cent) were not disturbed. Since most of the nests destroyed are broken up by predators it is evident that the concealing coloration of the bird and its habit of sitting close at the apjiroach of danger are much more eflective shields against danger than the thickness of the cover inmiediateK surrounding the nest. This is further borne out in the previous table. The Effect of Distance jroni an Opening:: on Nesting Success Investigation field men, early noticing a tendency of some predators. |)arti(iiiarly fox. to follow along the edge of openings and woods' roads, considered the possibility that nests lo- cated close thereto might be more subject to predation than were those located at greater distances. Table 133^ illustrates the results of this study. The percentage of nests hatching within ten feet of an opening was 53.6 per cent in comparison with 60.8 ])er cent for those located over 200 feet distant. Though both totals are based on a substantial number of nests, the dif- ference is not large enough to represent with certainty a valid trend though, in this situation, such a one would seem logical. In further subdividing the number of nests hatched at various distances into the cover types in which they were located, the numbers became too small to permit trends to be eval- uated except by combining similar types. This done, there still seems to be no substantial indication that nest mortalitv. at various distances from openings, varies markedly with the type of cover. Since nearly half of all nests are located along woods' roads or trails, a special study of such situations was made to determine whether or not nests are more likely to be broken uj) when located along such trails thati in the vicinity of other types of openings. As in- dicated in table 134^ of 359 nests along a woods" road or trail. 210 (58.5 per cent) were not molested while of 572 near all olhci t\pcs nf (ipcnings .'^13 (51.7 per cnii I haliluil. • See A)il>fni)ix, [•. 797. A See A|ii>fnlt -v:.<5C w^^^^'-^ji^^^^^r^ .,ft.,c -y.iit iirlr 138 COl'ER CHARACTERISTICS A\'n SHELTER REQUIREMENTS It is encouraging to find no important difference here, for woods' roads represent an in- tegral part of most woodlands which it is wise to maintain in the interests of providing more favorable nesting cover. The surprising thing about grouse nests is that so few cover influences seem to strongly af- fect them, either as regards location or fate. As pointed out elsewhere, the ruffed grouse has survived over a period of at least 25.000 years. Ample opportunity has thus been provided for the species, either to develop highly specialized habits and requirements or to become so adaptable as to fit into many different situations with reasonable success. Here is evidenre that the latter is the case. Brood Cover and Related Influences Turning from nests to broods, an even greater variety of questions arise. What types of cover most nearly meet their needs? How does this change with age, sex. seasons, weather, and with the time of day? What undergrowth conditions are to their liking? Are they par- tial to particular slope situations? These are but a few in need of answers. As long as one is discussing nesting cover, the ])robleni of recognizing grouse needs in terms of usage is not particularly complicated. It has been assumed, based on the collective expe- rience and judgment of the authors, that most types of cover were usually available to the fe- male in which to nest if she chose and that the best liked one was selected. Thus use became directly interpretable into choice. With broods and adults, the picture is scarcely as clear cut. Their requirements change with age, sex and the seasons. So also does the ability of the cover to satisfy them. A newly hatched chick, feeding largelv on insects and unable as vet to flv. is iiaturallv interested in a dif- ferent food and shelter combination than is a ten-week-old bird su])sisting on leafy material and fruits and quite able to take care of itself. An overgrown field may offer satisfactory refuge in summer yet fail completely to afford adequate shelter against winter weather. Other complicating factors exist. The quantity, quality and arrangement of cover are dif- ferent in each individual covert. This means that broods and adults mav not be able to exer- cise a free choice of cover types, undergrowth conditions and slope iiecause such are not equally available to all at the same time. Furthermore both quality and quanlit\ of cover t>pe var\ with tlic seasons and over the years. The latter may often pass by the unwary investigator unrecorded with tlic resuh that he, also, may not be able to exercise a free choice of cover type data. As a bird matures it gradually establishes its own territory, into which it. at times, resents intrusion by other grouse of the same sex. The jiest habitats may be fully occupied, thus forcing some birds to establish their tcrritoric-^ in the less desirable |)arls of a comtI. tliout;b not from choice. To make interpretation doubly dillicult. the iirobabilit) of obseiM'd brood or adult reac- tions not being typical is directly proportional to the disturbance of the normal response caused by the itucstigalor in the course of his work. One cannot design and carrv out a cotnprehcnsiM- study of the grouse and still fully com- pensate for the effects of all these complications. Their influence, however, may be minimized by securing a large number of records covering a iiNiiiiMr of \ears from earli of the repre- sentative grouse haliitats over the State. From thcsi' the significant trends usually can be deterinincd. This was the j)rocedure followed in ascertaining such items as cover preferences COVER REQUIREMENTS OF THE GROUSE 139 and the related effect of undergrowth type and density, weather conditions and slope on grouse. Individuals addicted to making snap judgments may fail to see the connection between a knowledge of brood preferences and an increase in the grouse crop. But those whose job it is to assist Nature in producing that crop, know better. As but one concrete example, grouse broods, in New York, at least, do not like steep slopes. A breeding area developed there is, then, going to be less productive than one developed on a gentle grade or where the land is relatively level. Many similar preferences, each small in itself but collectively important, must be ferreted out and catalogued if one is to go about the job of producing better grouse cover in an efficient, businesslike way. What, then, are the characteristics of good brood cover? Types of Cover Used by Grouse Broods The outstanding characteristics of brood cover are its diversity and its youth. It is the early stages of woodland succession, with their profusion of fresh herbaceous growth, that are attractive. (See table 13.51*. The abandoned pastures, the overgrown fields, the fresh, not too » Sop Afpcnilix. p. 801. ■iffZ,' ,^ Lr^rdmer bump THE FAVORITE HAUNT OF GROUSE BROODS THROUGHOUT THE SUMMER IS THE OVERGROWN FIELD, ESPECIALLY IF BUT FEW CONIFERS ARE PRESENT (TYPE c) 140 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS dense tangles that follow by a few years the cultinn; of the winter's fuel supply in the farmer's woodlot — these are the truly happy hunting grounds for grouse chicks. Of all the woodland types only second-growth hardwoods (type E) with its often abundant understory. and spot- lumbered areas (G) seem worth a second visit until the brood is well on the road to maturity. Where summer slopes are dry. as in our Adirondacks, the moister alder beds (B) are ])rime favorites. Least used are the t)pes where grass predominates (A), where the ground cover is rela- tively deficient, as in the shade of older woodlands I F. FH I or under a dense stand of conifers (HI. In fact, in New York, the j)resence of conifers, so dear to the hearts of grouse in the late fall and winter, is now almost ignored. Yet it is not easy to state definitely just which are the most used types of brood cover. There appears to be a rather strong relationship between the amount of cover and use. To be strictly accurate, one must reduce brood contacts to a "per acre" basis. It was not feasible to do this for all of the study areas from which the records collectively contributed to table 135*. Therefore, the amount of each type of cover down to one-tenth of an acre on the prin- cipal study area. Connecticut Hill, was determined and the number of broods recorded was adjusted accordingly as a basis for table 14. T.VBLE 11. TIl'KS OF COVER USED BY GROUSE HIU)()DS— CONNECTICUT HILL .\REA— 1930-1936. Type Type of code cover Number of years type was present on study area Average acreage of each type Number of broo<]s found in each type Adjusteds * Percent of broods found in eacli type Open liind Ovprprow n IiukI Woodlalui Slu»liin»: A H r. D Kll F FH C 11 I J 6.2 5.2 5.6 5.9 6.2 5.7 fiJ. 3.0 6.2 2.8 5.7 220.7 101.2 140.2 lll.l 445.8 469.5 159.0 46.7 58.3 134.6 63.9 1.36.7 14 78 148 65 I 10 28 5 3 31 32 39 65 .i 83 144 58 94 A .i .i 6.3 73 61 13.9 24.1 9.7 15.7 ibis 3.7 12.2 10.2 Total 58.3 2127.7 648 598 100.0 *The adjusted number of broods is the number to be expected if each cover had had the average acreage of 189.0 acres, and if each cover ty()e h«d been surveyed the average number of years. 5.1 years. Adjusted niiniber of hrnods ^ Y-O.HhO')\ ] -(1,1 I IO\'j; in which Y -= the observed number t)f broods iti a rover ty|>e, X j ^ the deviiitioM cif Ih*- r^uridnT of years ii coNer lypt- was survryt'd from the avmiL-e number of yeiirs surveyed per rover type; X2 = the deviiitio<|H observed in Cover Types A, I'MI, F and FK were so few tluit it was apparent that these ty|>es are h'ttle used by ^Toiise brtMHls. Outstanding among the differences noted is the elimination of second-growth hardwoods and conifers (EH) as an important hrood type and the increase in rating of s|)ot-hHnhered hard- woods (Gl. Unquestionahly. table 11 more accurately portrays actual hrood use. The com- parison still further serves to emphasize the desirability of oyergrowii hinds, young hard- woods and slashings as the types predominantly chosen by grouse broods. I hat second-grt)wlh hardwoods and conifers are neyertheless quite acceptable to broods has been evident on the Adirondack area whrit' thc\ are a more predominant clrmtiil of llic habitat. In this region, toit. aldci- runs ha\<' had a marked attra|irnilii. r'. «U>. t Srp A|i|trndii, p. BOS. COVER REQUIREMENTS OF THE GROUSE 143 ALDKK KLiNS ARE PAKTlCLi LAKL\ ATTHA(.TI\ K TO BROODS IN THK ADIRONDACKS SECOND-GROWTH HARDWOODS WITH A VARIED BUT NOT DENSE UNDERGROWTH ATTRACT MANY BROODS 1 H COVER CHARACTERISTICS AND SHELTER REQUIREMENTS glance a|)i)par to be significant, are probably accidental rather than valid, for a closer inspec- tion discloses no consistent tendencies. Most days, particularly in summer, are apt to be gusty or with moderate wind conditions and the number of broods contacted under either still or strong wind conditions are, therefore, comparatively small. Atmospheric Conditions. In comparing cover choice in sunny, cloudy and rainy weather, a rather interesting reversal of trends evidenced under conditions warmer or colder than normal, is observable (table 140)*. Apparently, woodland types ( E, EH, F and Hi furnish rainy weather shelter, whereas the more open overgrown and slashing types are less well patronized at this time. Conversely, in sunnv weather, the latter groups as well as the more open second-growth hardwoods, prove to be more attractive. Yet it is these same groups that are most sought out during colder weather, whereas, in warmer than normal weather, greater use is made of conifers, as has been previously mentioned. As in sunny weather, the open land types are apt also to be favored in cloudy weather. Ground Conditions. In attempting to plot brood cover choice under wet conditions, in con- trast with dry ground, the results are complicated by the fact thai, as soon as the youngsters can fly. they may seek shelter under conifers or in trees within the type. Thus, while the her- baceous cover may appear wet to the field man, the immediate situation where the birds are resting may be quite dry. One is prepared, therefore, to find from table 141'^ that grouse broods do not vary their choice of cover depending on whether the ground is wet or dry. The table indicates no strong tendency in this direction. Nor are the records consistent in this regard. Apparently, wet cover is no deterrent to the birds seeking food and rest in such open types as overgrown lands or slashings. Alder beds (type B) may then be used somewhat less, possibly because they are naturally a wet land type. Even here, the birds have been observed many times with their "feet almost in the water," for, especially in a dry season, some of the best food is there to be found. In retrospect, one cannot escape the conclusion that few weather influences seem to be sufficiently strong in themselves to govern the brood choice of cover types. Some Relationships of Undergrowth to Grouse Broods When the need arises, it is natural for most of us to classify a forested area according to the size and character of the tree growth. Only when the undergrowth contributes to the reproduction of the forest stand or is difficult In penetrate is it usually considered worthy of special mention. But grouse use dilTercnl standards, for. to tliem. both undergrowth and ground cover mav |)lay a substantial |)art in meeting their indi\idual food and shelter re- quirements. Just how iniporlatit that part is, has never been deterrnined. The Investigation, therefore, considered it wortliwhile to look into a few of ttie less cninpli'N influences of un- dergrowth on grouse distriliution. The undergrowth on tlie \arious sluiK areas was divided iiilu the fcillnuing six types: — 1. Large herbs 2. Shrubs and berry bushes 3. Hardwoods 4. Hardwoods and conifers 5. Conifers 6. Combination of lv|ies ♦ Set Appcnitix. I'. 806. A Sec Appcn. 812. t See Appendix, p. 813. COVER REQUIREMENTS OF THE GROUSE 149 As on the Connecticut Hill area, from this it would appear that steep slopes are markedly less attractive to broods than are level lands or moderate inclines. It will be remembered that few nests were found on abrupt hillsides. Perhaps the reason for this is that fewer grouse live there. Be that as it may, both observations point to the desirability of including as much good grouse cover on the less rugged slopes as possible in any lands on which a large partridge crop is to be encouraged. That few sportsmen are hardy enough to hunt there, and the likelihood of encouraging severe soil erosion should one attempt to improve the existing cover by judicious cutting, should also not be overlooked. There are a few minor variations in the choice of slopes at various ages to be found in the table. A majority of the broods at all ages (63.9 per cent) are found on gentle slopes. Steep hillsides are used very occasionally either by the very young birds (probably recently hatched I or by the broods as they approach maturity. In the Adirondacks flat alder beds are used to a pronounced degree throughout the brood period, although less during the latter part of Aug- ust than earlier. Slopes Used by Grouse Broods at Dijjcreiil Times of Day After considering so many of the forces which might, but seem not to. strongly affect brood cover choice, it is reassuring tu find the chicks actually exhibiting, in table 148*. apparently valid preferences for certain slopes at certain times of day. The number of broods found on a certain slope during an) particular hour ma\ at lirsl glance be misleading for the records are so few as to vary considerably. By glancing at the graph*, however, the eye can pick up trends which are interesting though difTicult to explain. No slopes are really shunned at any particular jieriod. There is a trend for more grouse broods to frequent the cooler north slopes in the rnorning rather than in the afternoon. Likewise the east slopes, open to the morning sun. a|)pear to be more attractive eai I\ in the day and again in the afternoon rather than through the late morning hours or at iniil(la\. It is then that the warmer south slopes attract more broods. No particular preference for western exposures at any special time of day is observed. Cov- erts without slope, that is flat lands, are likewise about cqualK well patronized throughout the day. As with many another link in the chain of life-history facts about the grouse, it is far easier to chart the trends than to offer logical explanations for the underlying reasons therefor. It seems evident, however, that no exposure is so uncomfortable at any certain time of day as to cause broods definitely to avoid it. Some Relationships of Temperature, W iiid and Atmospheric Conditions to Slope Use It is often productive to explore even the seemingly inconsequential relationships between a wildlife species and the habitat which nurtures it. Like a complicated puzzle, each piece of information that can be fitted into its proper niche helps to interpret the whole even though individually it may be of but little immediate moment. The differences in use grouse broods make of cover facing north, east, south or west under varying conditions of temperature, wind velocity and moisture conditions, are apt cases to point. For instance, if the birds were disinclined to make use of otherwise satisfactory cover because it was located on a slope facing strong prevailing winds, such situations might be * See Appendix, p. 81 1. 150 COVER CHARACTERISTICS A.\D SHELTER REQUIREMENTS among the less desirable for inclusion or development in a grouse management area. The only sure way to answer such questions is to check carefully each such situation by way of finding out just how important it may really be. In considering the effect, complicated as it is, of these influences upon grouse, one must remember that such items as the way a slope faces and the temperature conditions found there- on, certainly are responsible in some measure for the vegetation which occurs. For instance, in many parts of New York the colder north slojjes run to the beech, birch, maple, hemlock type. The warmer south slopes are often covered with types such as oak and cherry which are common farther south. One generally thinks of animals as seeking south slopes for warmth during cold weather and north slopes for relief from the sun during warm periods. In table 149* we find that grouse broods seem to react in a reverse manner. The use of north, east and west slopes is appar- ently least on warm, greatest on cold days. Conversely, south slopes and flat lands are occu- pied least on cold days, though the difference is admittedly not great. Thus, while this table does show interesting brood choice trends under various temperature conditions, it is probable that cover types found on the slopes, rather than the exposure of the slope itself, may be largely responsible therefor. Produce the proper admixture of cover, and slope aspect may not be important. To do this where it does not exist, may be a costly procedure. It is well to remember that, in New York, tracts containing large areas lying all on one slope may be less easily made into high producing grouse units than those in which a natural variation in tojjography may assist materiallv in producing naturally the variety of cover grouse like. Wind, too, may have its effect on the degree to which grouse broods frequent various slopes. While none is shunned by broods primarily because of wind conditions (table 150)^ there is an indication that the birds keej) away from north and west slopes on windy days. This is logical when one remembers that prevailing winds over much of New York are from the west, and to a lesser extent from the north. Further developing this impression, the table indicates that east slopes and flat lands may be somewhat more attractive to grouse broods when strong winds are prevalent. Unfortunately, no record of wind nings. Cover use varies with the time of day. though no type is com|)letcly shunned at any time. Weather and slope influence brood distribution dej)ending on the makmip of thi' crown cover and undergrowth but the effect is seldom strong. Broods lend to ;i\iiid steep slopes. • Spr A[ipcnilix. p. fllfi. A Srp Apprntlix. |«. 816. t Spc Aiiprndix, p. 817. COVER REQUIREMENTS OF THE GROUSE 151 Adult Grouse Cover and Related Influences It is a pleasant part of every day's hunt to talk over in retrospect the experience of the chase. Here a grouse was found feeding on frosted fruit under a wild thornapple tree. There two birds flushed unexpectedly from a thicket of alder or birch. Conifers screened a third that got safely away. Where were the rest of the partridge, and why were they, too, not found? To the old-timer each new day's adventures, mellowed by experience, furnish fresh evidence to substantiate impressions as to the probable behavior of the bird under a wide variety of con- ditions. In a practical sort of way he becomes his own "expert"' in predicting where they are most likely to be found and where they will fly to when flushed. His judgment may not always be vindicated, for each grouse is an individualist in its own right, but there is never- theless much of truth in what he may say on the subject. Wildlife managers, listening in, may perhaps be confused by the variety of often conflicting opinions thus expressed. But to date there are few places, save for their own limited expe- rience, to which they can turn for more exact impressions. As we have seen, in attempting to chart brood preferences for certain kinds of cover, slope, weather and wind conditions, it is no easy job to interpret correctly grouse behavior. The background of environmental influences is so varied as to require the most careful examination of thousands of records be- fore one can even recognize a possibly valid tendency for the bird to ad in a given manner under a certain set of conditions. Yet there is no other way, known at the present time to an- alyze grouse cover rpquiremcnts with sulTicient accuracy to justify the expenditure of hard cash in improving the habitat to pnxhicc a larger grouse croj). In no other way can we sort out the ideas and determine which of the old-timers' theories have real substance in fact. Let us then look at the record written by the birds themselves. To establish a pattern in which we can place confidence, individual variation has been largely compensated for by an- alyzing 19.619 records of adult flushes from tlie three main regions of the State. To leaven out any radical differences which might be apparent in any one year all flushes from 19.30 through 1936 were tabulated. To insure representation from each region, four study areas totaling 10.000 acres were set up. one each in the Adirondacks and the (^atskiils and two in the Southern Tier counties. Adult grouse contacts on these areas alone totaled 16.963 dur- ing these years. In charting grouse reactions tt) some items such as cover types and slope still furliier re- linenients were necessarv. The amount of each type and slope had to be determined and the record of birds found in each situation adjusted accordingly. If. on a study area, one type of cover occupied much more of the land surface than another, yet both were equally attrac- tive, birds might be expected to be found thereon in proportion to the size of each area. This would result in a seeming prei)onderaiice of birds being found in the more extensive types unless su(ii aTi adjustment were made. One ("mds ibis difference evident in comparing tables l.S and 1.^3*. To correct this situation the number of flushes in each type of cover had to be expressed on a "per acre" basis. In making up some tables it was even found necessary to consider the changes in the extent of each cover type taking place while the Investigation was in prog- ress. Some of these were surprisingly large as abandoned meadows grew up to brush or small slashings were cut only to fill in to sprouts and berries in a few years. To keep track of such items on all the study areas was a task beyond the resources at * See Appendix, p. 819. 152 COVER CHARACTERISTICS AND SHELTER REQUIREMENTS hand. The Connecticut Hill area was; therefore selected as the one on which these changes could best be noted. Thus the following tables from which the pattern of adult grouse be- havior here described, is drawn, may represent one area, all four, or records from the entire State depending upon the complexity of the situation to be analyzed. No exact way of taking into account differences in the arrangement or interspersion of cover types was found. Realizing the im|)ortance of having each type well scattered over an area selected for study, only units supporting a good admixture of cover were chosen. To the extent to which each varied in this respect (and others such as the quantity and quality of the vegetation making up a cover type) inaccuracies may have crept into the tables to influence the trends here presented. In sampling the effect of the environment on a species such as grouse, no way of compensating for these has been discovered. Fortunately they are probably not too large in most instances to obscure seriously the more pronounced trends. With this understood, let us see what evidences of grouse reactions to cover have been fiiiind. Types of Cover Used by Adult Grouse Grouse broods love the lush confusion of slashing and brushlot. As the birds mature and the broods disperse, the approaching winter poses new problems of finding food and shelter. The overgrown lands and brier patches rid longer provide the same luxuriant protection once the leaves have fallen. Only the conifers and the broad-leaved evergreens maintain their prom- ise of sanctuary against winter winds and winged predators alike. Tangles of vines too dense to be attractive while the leaves were on. now, with a mantle of snow, furnish places to doze in fair security. As winter gives way to spring, and spring to suinnier. cover use reflects the vegetative changes in the face of field and woodlot that accompany the seasons. The habits of adults are modified accordingly. Remembering the brood cover preferences, one nia\ be surprised to find indicated in table TABLE \?>. TYPES OF COVEK USED BY ADULT c;HOUSE>— CONNECTICUT HILL AREA— 1930-1936. Type had been surveyed the avorafcc number of yearit, 5.1 ycarti. Adjusted number of adults = Y — 0,8609Xl — O.IMOXi': in whirh Y = the tibm-rved number of aduttn in a rover type. Xi — llie deviation of th^ niinibrr of years a cover type Willi Nurvcvetl from thr avcrAge number of years surveyed per rover type; Xs =: the deviation of ruiiiihcr of acres in a cover lv|»r- from the averace number of acres per cover type; 0.6609 = average increase in number of adult?* with each year increase in time» surveyed; 0.11(0 '=■ averace increase in number of adults with each acre increase in cover type. A The number of adults observed in cover type A was so few thai it was apparent that this type is little used by (rouse adults. COVER REQUIREMENTS OF THE GROUSE 153 15 a generally lessened interest on the part of the aduh in overgrown lands and in slash- ings* though many a bird still seeks the abundance of food to be found there, particularly during the summer and fall months. Second-growth woodlands, though usually well patron- ized, are also less important, unless they contain at least a scattering of evergreens. Con- versely, the adults seem to find the more open ranks of the mature hardwoods and conifers, as well as the denser stands of evergreens, much more to their liking than did the broods. In reducing grouse flushes to a "per acre" basis, one source of error has been avoided, but another encountered. The scarce types, even though moderately used, have been made to ap- pear more important than would actually be the case, were they, in fact, more extensive. In other words, use is not proportionate to size. In using a "per acre" analysis, types large in area are made to appear less important whereas types represented by but few acres appear to support a larger use than is actually the case. This situation should be kept in mind when interpreting the cover preferences by month on page 157. a.s well as in other tables showing "birds per acre." To be of much use the general cover picture must bo broken down by seasons and months. Let us then take a look at where the adults are then to be found. Seasonal and Monthly Cover Preferences Among the questions frequently asked by sportsmen during the course of the Investigation has been, "Where can I find the most grouse?" It is usually motivated by the desire to har- vest a share of the rriiifcr> Spot-lumbered areas iG) Relatively pure stands of iKipp Younger slashings (I) IHl l.inli alder (Hi COVER REQUIREMENTS OF THE GROUSE 159 Month Types most jrcquently used (cont'd) Conifers (H) December Mature hardwoods and conifers (FH) Second-growth hardwoods and conifers (EH) Older slashings (J J From the above, one might well gather that grouse in New York State cannot exist where conifers are absent. Such is not the case. True, they are an outstanding part of most of our productive coverts. But when they are absent, other types, such as spot-lumbered areas (G) and older slashings, may furnish brushy tangles which serve the same purpose in pro- viding escape cover and shelter. In general, however, the more severe the weather, the more conifers, properly interspersed, contribute to the desirability of any grouse habitat. The above "design of living quarters" for adult grouse month by month is of course made up from the observed preferences of the birds under New York conditions. By and large, prob- ably it represents a fair cross-section of grouse cover choice throughout the Northeast. Types of Cover Used by Adult Grouse at Various Times of Day In tables 155 to 158*, one has, in effect, put a time clock on grouse movements from cover to cover throughout the day. Hunters, bird lovers and research workers alike are interested in finding out how these birds budget their time. This is no easy task, for a quick glance at the tables leaves one jiuzzled by the complexi- ties of the use pattern to be unraveled. In truth much of it cannot be assembled, for few- strong trends stand out even when the records are broken down, season by season. In other words, there is no outstanding coticentration of birds in any one type at any particular time of day. But there are some interesting seasonal use differences to be noted. Winter. The necessity of finding food and sheUer seems to be the controlling influence on grouse movements throughout the winter day (table 155) . In the early morning, in general, the birds are most likely to be found among the conifers (type Hi or the second-growth hard- woods (E). The latter type often combines good budding opportunities with a chance to catch the morning sun. While overgrown lands and slashings are used least of all in winter. one finds a few early feeding birds in the former covers at this time. The most unexpected thing about early morning use is a tendency to avoid second-growth hardwoods admixed with conifers (EH). The table clearly indicates that birds do not move into this type in numbers until later in the day. By mid-morning less birds are to be found in second-growth hardwoods alone; more in a mixture of these with conifers. Little change is noted in the attractiveness of overgrown land and coniferous patches. At mid-day mixed hardwoods and conifers reach their maximum use. Birds now tend to forsake the alders, popples or birch thickets (B) and to be somewhat more prevalent in the older cut-over areas (J). Many still stick to the conifers though possibly in less numbers than is the case earlier in the day. Towards late afternoon more birds may be found feeding in the overgrown lands. Sec- ond-growth hardwoods and conifers (EH) likewise are attractive but relatively pure stands • See Appendix, p. 823 to 829. 160 COJ'FR CHARACTERISTICS AM) SHELTER REOl IREMENTS of conifers iHl are likeh In lia\e less liirds aiiKitij; llieiii llian at aiiv otliei time. Spring. The daily cover use |)att(rn in spring is markedly different from that of the winter ftable 156)*. As one would expect, nesting cover (types E. EH) is now important. Here one is apt to find about the same percentage of birds throughout the day. Conifers (Hi. while less used now than in winter, are still frequented from daylight to dark. In early morning a relatively large number of adults are to be found in the more open overgrown lands iB and Cl except where young conifers predominate (D). Possibly thev are in search of variety in their diet. Interestingly enough this tendency does not extend to cut-over areas (I and Jl. however, since these are but little used until later in the dav. By mid-morning this situation is apt to be reversed. The more open "edge" types (A, B and C) appear to be utilized less, except for overgrown lands dotted with small conifers (type D), which for some reason, now gain markedly in popularity. Some birds also seek out the older slashings (J). From midday through late afternoon this pattern changes but little. As in winter, alder, popple and birch cover (B) are progressively less used; brushy areas (C) more. Possibly a few more birds may be found among the conifers, but the shifts are, at best, minor. Summer. Two differences from other seasons stand out in summer cover use (table 157)^. Thick, coniferous covers are but little frequented. Even mixed second-growth hardwoods and conifers (type EH) are much less popular than is hardwood cover alone lE). There is also a strong tendency to make greater use of shrubby and herbaceous areas such as brushy pas- tiires, spot-lumbered areas and slashings ( B, C. D. G. I and J I. Here birds find summer feed in unending variety and abundance. Early on a summer morning birds are apt to be found in the overgrown fields (types B. C and D) or in the second-growth hardwoods (type E). In fact the latter type is popular throughout the day. Around midday the older slashings (J) seem to be particularly attractive, though the cover just mentioned is also frequented. Late afternoon finds the birds more widely distributed though possibly with a tendency to work back into second-growth hardwoods (E). Eall. In some respects good fall grouse cover is not as different from that used by the birds in summer as one might think (table 158)*. As every grouse hunter knows, birds are to be found in the overgrow'n lands (types B, C and D) in considerable numbers. Hedge- rows and brushy patches, not too far from wooded areas, are favorite gunning spots in early fall. But unless it is a "beechnut year" one has only about half as much chance of finding birds among the second-growth hardwoods (E) as in the sunnner. To add a fair sprinkling of conifers, among the young hardwoods (EH), however, is to increase the popularity of this type considerably. Small ])atches of conifers (H) are three times as likely to be used in au- tunm as during the warmer weather. Even so the birds are apt to be sur])risingly well dis- tributed over most types, the notable exceptions being open land (Al and mature hardwoods (F). Apparentlv. too. cut-ov<'r areas (I and J) serve as sunnner and early fall feeding grounds for later they attract somewhat less birds, except in the early morning. Curiously enough. on the Connecticut Hill area, young slashings (I) were also much frequented in November. • Sec Apiirndix, |i. 62't. A Sec Appendix, p. 827. t Sec App.-ii.lix. p. «-"'■ COVER REQUIREMENTS OF THE GROUSE 161 Sunup is the time to hunt the brushy spots and overgrown pastures particularly where there is a dearth of small evergreens (C). Birds are apt to be found also in mixed second-growth hardwoods and conifers (type EH) and in clumps of conifers (Hi. A considerable propor- tion pay an early morning visit to the older cut-over areas ( J I . By mid-morning there is a tendency for birds to move back into the wooded areas, either into the two woodland types mentioned above or to second-growth hardwoods (E). Midday is apt to bring on a small increase in birds among the popples, birches or alders (B) and in mature hardwoods with (FH) or without conifers (F). There are also likely to be fewer birds among the conifers (H) even though they be in mixture with second- growth hardwoods (EH). No significant changes from this pattern are indicated throughout mid-afternoon except that conifers I H ) are still less popular. This is apparently the best time of day to hunt the spot-lumbered ])atches (G) especially if they are not too extensive. The older slashings also show a small increase in use at this time. It is interesting to find no great increase in the numbers of grouse frequenting overgrown lands (B, C and D) and but few birds in cut-over types 1,1 and J I in the late afternoon in spite of the widely accepted belief that such coverts are much used for feeding at this time. Woodland coverts containing a mixture of conifers (EH and H). however, exhibit in- creased use. Thinking that tin' cover preferences of grouse dining llic hunting season (October, No- vember) might differ significantly fmrn the overall fall picture, flush records gathered from all parts of the Slate for these months, were a.isembled in table 159*. An analysis of these, however, reveals no great difTercnce between the cover preferences of the birds in September when compared with that of the later autunm months except that, as the days grow colder, less use is likely to be made of cut-over areas (I, J) in the early morning. Effect of Various Weather Conditions on llir Choice of Cover by Adult Grouse The reaction of grouse to differcril weather conditions is naturally of interest to the hunter. Such relationships may also he important to the game manager in i)reparing development plans. Accordingly, the temperature, wind and atmospheric conditions prevailing at the time each grouse was flushed, were noted throughout the Investigation'^. As a general conclusion it a])pears that grouse may be found in any cover type under any of the weather conditions recorded. However, the types predominantly utilized at any one season tend to outrank the others regardless of the weather. But an analysis of tables 160- 162^^ does reveal interesting differences of cover use under various weather conditions. Temperature. Considering temperature first, the data have been summarized in table 160. At a glance, an inclination for the birds to move into mixed second-growth hardwoods and conifers (EH) or to thick conifers (H) on colder than normal days is apparent. This seems, however, not to extend to mature mixed hardwoods (FH) or to the more open over- grown lands with an interspersion of evergreens (D). This suggests that the higher shelter value of the denser woodland types may be the deciding factor. Thus conifers may serve the adults as protection from cold in winter but not from heat in summer. Conversely, on warmer than normal days, grouse tend to move to some of the more o])en * See Appendix, p. B.'iO. A See figures 73 to 76. p. 702. t See Appendix, p. 831 lo p. 833. 162 COl EK CHARACTERISTICS AM) SHELTER REQIIREME.XTS rover typfs deficient in eonifei>. Anions; these are alders. ]K)|)]de or Lireli i)atclies (Bl, sec- ond-growth hardwoods (E), mature hardwoods (F) and >()ung slashings I I). This rehi- lioiiship does not hold true, however, for overgrown fields, with (C) or withoul iDi an ad- mixture (if conifers. No good reason for this seeming contradiction is apparent. In considering the above, one should bear in mind that the designations warm, normal and cold refer to the relative temperatures at the time a flush was recorded as compared with the average for that time of year rather than to the actual temperature in degrees. Wind. It is generally believed that, on windy days, grouse often exhibit a tendency to "flush wild." Certainly, in windy weather, they are more alert and nerxous than at other times. One might exj)ect to find, and be surprised at not finding, upon examining table 161*. the birds concentrating largely in shelter types at this time. True, coniferous clumps (H) are then more likely to be used but mixed hardwoods and coniferous cover (D. EH and FH) exhibit no such tendency. The only other covers, more attractive in windy weather, are the alder runs or stands of pop- ple or birch (B). Particularly in the case of alder runs, this seems related to the generallv protected locations in which they usually occur. Possibly there is also a tendency for simi- larly situated portions of other types to be preferred at such times but it has not been feasible to record the data covering this point in such great detail. The table also indicates that more birds frequent second-growth hardwoods (E) on wind- less days than at other times, though the reason is not clear. Thick coniferous cover (H) appears less likely to be frequented. Other types show no significant differences in use in windy versus still weather, all of which leads one to surmise that wind normally exerts a rather minor influence in grouse cover choice. There is, of course, the possibility that, had the data been broken down according to the season of the year, greater differences might have been found. Atmospheric C.oiuUlions. In general, sunshine, rain or snow seem not to exert any pro- nounced influence on the general pattern of grouse cover choice (table 162)" except when shelter is needed. During snowstorms the birds, as expected, are most likely to seek out heavy coniferous cover iH). though, as with wind, this tendency does not extend to mixed hardwood and coniferous cover ( D. EH and FHl. Likewise the more o|jen cover, even though some conifers are present, shows a decline in use at this lime. Rallii'r curi- ously none is completely shunned even in snowy weather. Rain apparently bothers grouse less than snow. Thus one finds at this time fewer birds resorting to the more adequate protection afforded by thick coniferous stands (H); more birds preferring the mixed second-growth hardwoods and coniferous cover (EH). There is also a curious tendency for open fields (A) to be frequented more on rainy days. \^'hile worth noting, this is of no real im])ortance since little use is made of this type at any time. Several of ihc more open types are used more in sumiv weather than at anv other time. This is true of overgrown fields, lacking in conifers (C). spot-lumbered areas (G), mature hardwoods (F) and the older slashings (J). Conversely, heavy coniferous stands (H) are tiien least attractive. Effect oj Various Ground Condi/ions on the Clioicc of Cover h\ Adull Crouse Ground conditions exercise a greater effect on grouse cover choice than is generally realized * Sdo Appendix, p. 832. A See Appendix, p. 633. COVER REQUIREMENTS OE THE GROUSE 163 for there is considerable variation in type use depending on whether the ground is dry, wet or snow-covered (table 163)*. For instance, we have already noted that conifers are most at- tractive in winter. This is particularly apparent when snow covers the ground, for 40 per lent of all adults sought this type (Hi when the snow was moderately deep. Conversely, the more open types in which no. or but few, conifers are present are less used at this time. An interesting sidelight is the tendency for some birds to abandon conifers for relatively open, mature hardwoods (F) when deep snow encourages snow roosting. Cover use trends are less predictable when the ground is dry, damp or wet. Overgrown lands deficient in conifers (C) and the older slashings f J) seem to be most attractive during dry or damp periods, with the birds showing increased interest in coniferous stands as the ground becomes wet. Second-growth hardwoods lE) and mixed hardwood and coni- ferous cover, irrespective of age (D, EH and FH) are. however, frequented to about the same degree, irrespective of these conditions, thus raising the question as to Imu much these differences really matter. Preference of Adult Crouse for Ground Over Trees Like other gallinaceous species, the grouse spends most of its time on the ground at all seasons of the year, as shown in table 161'^. \c\frtlic]ess it often resorts to trees, especially conifers, for roosting, or when disturbed by a predator or some other inlruilcr. Budding in i.irfitnrr Bump WHILE PREFERRING THE C-R3UND. GROUSE TAKE READILY TO TREES WHEN NECESSARY trees is connnuii during the colder months. Tree roosting has been most frequent during the winter and spring, with only an occasional bird to be found therein during the other seasons. Strong winds further discourage this practice. * Sec App<"ni1ix. p. 83-1. " See Appendix, p. 8.'?6. 164 COVER CHARACTERISTICS A\D SHELTER REQUREME.XTS Table 165* was prepared to determine if this preference for the pniuiid varied according to the type of cover. It is evident that this habit has held true in all t\ ])es. \^'hen tree roost- ing has been resorted to, coniferous stands (type H) and mixed hardwoods and conifers (EH and FH) have been most frequented because evergreen trees are usually preferred for this purpose. Considering the few trees normally present in open lands (A), overgrown fields (B. C and D) and in slashings (I and J) it is surprising to find that from 4.7 to 10.8 per cent of the birds found therein were flushed from trees. Perhaps this is a reflection of the tendency for some trees thus situated to fruit abundaiitlv and for the birds to seek these out in the fall. Influence of Slope on Adult Grouse Distribution Had it been shown by a study of table 166'^ that grouse are more likely to frequent one slope than another at any particular time of day. birds might be more easily located. W itli but one exception no such inclination is found. At midday a quarter more birds were flushed from north slopes than at any other time. It is at this period that manv birds prefer to rest quietly in some sheltered spot. A'orth slopes in general are rather more likelv to support conifers, in the area studied, than are f)ther aspects. Evergreens furnish acceptable shelter. Herein, possibly, lies an explanation. A question also arises as to whether or not temperature exercises an appreciable effect on the number of birds frequenting any particular slope. Table 167' indicates that no aspect of slope is avoided whether the temperature is warmer than usual, normal or colder. Though the trend is not pronounced, there is an inclination to use west slo])es more and east less, on days which are warmer than normal. Likewise north and south slopes are less used when the weather is colder than normal. At this time flat lands seem to be most attrac- tive. The prevailing winds are from the west «n the areas studied. The underlying reasons for such a|)pareMt ])iefcrences arc not always easy to fathom. Yet all these observations are statisti(all\ signifuanl. In other words the chances are at least I'J Ic) 1 that these conclusions lune a sound basis in fact. Thus the greater use of east slopes and flat lands in colder than normal weather points to the dcsirabilit\ of encouraging winter shelter co\er in such situations, insofar as is practical, when drawing uj) plans for improving grouse coverts. Summing up, good adult cover is characterized by a variety of composition and a high interspersion of cover types. Woodlands, particularly if they contain some conifers, are the backboni- of the covert. Overgrown lands and slashings furnish important summer and fall feeding opportunities. Though no types of cover are completely shuimcd at anv season, inmith. or time of day. there exists a fairly definite time-use pattern. Temperature, wind and atnH)spheric conditions exert a inodilxing influence on cover choice. In most instances, the differences thus caused arc not pi unced. Ground conditions, partic- ularly snow, also are responsible for some shifts in cover use. Slope exerts a minor influence. By studying the details of the points here presented, then- emerges a generalized '"design of living (]uarters." which, when |)roperly put together >lioulil fulfill all grouse co\er re- * See Appriiilix, p. 8.17. A See Ap)>cn«tix. p. 838. t See Appriidii. p. 8.19. t Sec the Bocliiiii »>n "Tlif Holi- of (^ivi-r Ciiiiipitnilii>ii .mil Vrriiiiccinfiir*. p. I(i8: iiNu llii- rlidpli-r^ . This is one of the reasons why large blocks of coniferous reforestation are apt to be productive of grouse largely along their edges*. The age of the trees or shrubs that make up a type also exerts an influence on the cover. Uneven-aged stands create breaks in the crown canopy. These let more sunlight filter through thus encouraging a more abundant variety of shrubs and herbs beneath. The effect of removing a few trees here and there in a woodland, known as selective cutting, has been noted consistently in terms of increased use of the remaining cover (type G in the foregoing tables) by botii broods and adults. The physical condition of the soil also has its effect. Fewer species are likel\ to thrive on lands from which the fertility has been exhausted. Conversely, the presence of a diver- sity of soils running from acid to alkaline, from wet to dry. and from gravel to loam, pro- vides encouragement for a greater variety of plants, many of which are adapted only to par- ticular situations. Such conditions, though seemingly small, should be taken into considera- tion by an individual or state interested in purchasing or developing areas for grouse. On previous pages, the impression may have been created that, as long as grouse rover contains an abundance of the various types which grouse frequent, it will produce a high population of birds. Such is certainly not ahva\s the case. Patches of almost pure conifers, dense hardwood sprouts, hawthorn thickets and second-growth maples, lying close by one another, still would not provide acce])table winter shelter, spring breeding grounds and summer and fall feeding areas to sui)|)ort many grouse. To be really productive there must be present not nnlv a good arrangement i)f((i\er t\pes but also a variety of vegetation within each. Of the two ideas, the former is. in principle, the belter understood. One thinks of a badly laid out farm as a poor producing unit. In like manner grouse cover tliat meets the food and shelter requirements of the birds for only a part of the year cannot produce a good crop of partridges. The individual cover types, like the separate farm fields, may partially fulfill the need of bird or farmer as the case may be. yet the whole be so organized as to spell failure of the year's crop. The analogy has been used purposeK. though maii\ sportsmen as yet do not realize that iheir shooting in the fall is largely dependent upon the co\er being so i)Ut together as to sat- isfy the birds. Fortunately, in New York Stale, the axe and the economic conditions creating much little used or abandoned farmlands have accidenlalh produced man\ fine coverts with a high carrying caiiacily for grouse. The\ are the ones In which the hunter returns year after year, always to find some birds. Hut there are others, and lhe\ mc in the majority, which are less frequenlh hunted simply because ihe size of ibe bird crop is less predictable. The most likely cause of this is poor arrangement of types. \o winter wood has been cut there for some years; no cows have tramped down and ihtis kept the brushy pastures semi- open. Almost imperceptibly the trees are closing in and growing taller. The light-demanding vegetation is being gradually choked out. Here year by year individual grouse territories are becoming larger and birds fewer as second-growth and mature cover types extend their boundaries and the edges retreat farther and farther from their centers. Realizing this, wildlife managers responsible for developing better grouse coverts are plamiing their plantings of trees and shrubs in abandoned fields to produce the maximum of edges compatible with fulfilling other uses. Large blocks of conifers are giving way to strip » Scr tahlc IB. [<. 171. 170 COVER CHARACTERISTICS A.\D SHELTER REQUIREMENTS plantings interspersed with hardwoods. Where practical, a greater variety of species is being used. The rich vegetation characteristic of many field edges and hedgerows is encouraged. Cuttings in existing woodlands are so planned as to break up large areas of continuous forest cover. All these are moves in the right direction, but their influence is. unfortunately, not as imme- diate or widespread as one might visualize. A year may produce a few oak leaves on a seedling planted in a recently abandoned field, but at least a generation of oaks is required to restore to the impoverished soil a fertility sufficient to support the woods' sedges and ferns, the wintergreen, partridge-berry, and the profusion of other herbs on which grouse feed at one time or another during the year. True, the pines may furnish shelter before they are head- high, the cherries and apples some food soon thereafter, but a rich profusion of woodland undergrowth and ground cover takes time to establish. Thus it becomes increasingly impor- tant on such lands that one encourages a diversification of vegetation by planting the proper (.KOUSK IIIKIVK WHKKK TUKKt: IS A RICH 1)1VKKSIT\ OK T\ I'tS. HERE OPEN FIELDS. OVERGROWN LAND, SECOND-GROWTH HARDWOODS AND MIXED HARDWOODS AND CONIFERS COMBINE TO MAKE A PRODUCTIVE COVERT species in the proper relationshij), one to the other. Likewise one should encourage those farming operations and wood-cutting activities that tend to maintain a ricli diversity of types. Suggestions covering this are to be found in the succeeding chapters on food and on management. THK VALUF OF EDGES In discussing the makeup of cover for grouse we are venturing into a field long traversed by woodsmen but little known to science. When this study first started, the catchy idea that to plant a block of trees was to increase cover for wildlife, was still in the sales manual of many a forester. It didn't matter much what kind of trees were set out. how far apart they were, or where they were [)ut. It was Leopold who first pointed out in 1936 that "the ])oteMtial density of game of low cruising radius (such as the ruffed grouse) is, within ordi- nary limits, proportional to the sum of the type edges." Today the need for diverse makeu)i and interspersion of types is itetter understood by both forester and game manager. THE VALUE OF EDGES 171 Effect of Edges on Adult Grouse Distribution In the old pastures and farm woodlots which characterized three of the four study areas, the axe and the plow had been quite successful in breaking down the original forest into an infinite variety of cover. Faced with the problem of classifying the vegetative groups, the Investigation recognized 12 major types as heretofore described. To determine the "edge eflect" of each type on grouse distribution, the location of each adult, brood and nest found over the 13 years of the study was plotted as accurately as possible on large scale maps. A copy of a single section, chosen at random but representing adult flushes for all seasons of the year, is here reproduced (figure 14). No strong tendency for the birds to be found along cover type edges is to be observed on many of these maps. figure 14. distribution of adult grouse flushes on one compart- ment of the CONNECTICUT HILL study area — ALL SEASONS, 1932- 1940. EACH DOT ON THE MAP REP- RESENTS THREE FLUSHES OPEN HAROWOOO HARDWOOD B MUDWOOOS HftROWOODS BflUSH CCniFCn BRUSH 3*- <2* B CONIFERS A further test was then employed. On the cover type maps of 12 sections of the Connect- icut Hill study area, a series of 20 random lines* were run. From the number of type edges crossed by each line an index of edge frequency for each section was obtained. These, tabulated with the corresponding average spring population densities for the period 1930- 1942, are indicated in table 16. Studying this, one finds that, in general, the larger the number of edges, the higher the average population of grouse the areas will support. * The posilion and direction of each line was eBtablished by Btatisticallv valid random sampling methods. 172 COVER CHARACTERISTICS AM) SHELTER REQL IREME.\TS .^ TABLE 16. MK[ATI().\SH1I> OF THK KMIXHJKNC.V NMTII WHICH KDGliS OCC.UH TO THE WKR.VGK LKVKL OF S1>1UN(; POPULATIONS ON 12 SECTIONS OK THE CONNECTICUT HILL AREA— 1930-1942 Avprapr diMisity of sprinf; Section Edge frequency |K>|inlutions per lUU acres index of cover 15 1.7 6.5 9 2.0 6.1 5 and 6* *> I 4.9 .IN 2.2 9.2 1 2.3 5.5 0 2.1 T.:i 7 2.6 10.» i 2.7 8.7 :is 3.0 7.8 11 3.1 8.7 K 3.3 8.6 * These Iwo sections have been ennsidered together sinre the onK line oi deiiiarkatinii between them is a small stteam. Effect of Edges on Brood Distribution It has been the general impression that broods, like adults, are inclined to seek out the more varied edge vegetation. Table 152* indicates the number of broods of different ages contacted on the Connecticut Hill area within 100 feet of the edge of one type of cover or a combination of two, three or four type edges. From it one gathers the impression that, irre- spective of age, the broods do not particularly frequent the immediate vicinity of several type borders. Realizing that the chance of finding the birds near where two tyi)es border one another is considerably greater than where one type corners on two or three others, the relative degree of probability for each of the four situations was determined by selecting from this area 100 points at random"^ and listing the number of t\pes to be found within 100 feet of each position. The results are shown in table 17. TABLE 17. HKL\T10NSHIP OF THE NUMBER OF COVER TYPES WITHI.N 100 FEET OF BROODS CONTACTED TO THE PROBXHILITY OF CONTACT AS DETERMINED BY RANDOM SAMPLING- CONNECTICUT HILL AREA two- ly 12 Number of crown cover types within 100 feet of cdiitjict point Percent of random points in each group Pereent of broods located in each group 1 i:i 51 2 12 38 3 12 9 4 3 2 The percentages here given for each group are so close as to ifiid fmllici weight In the cuiiclii- sions that broods do not particularly seek out situations where three or four type edges are to be found. Relation of Edges to Nest Location Other factors being equal, the desirability of a nesting site varies inversely with its distance from an opening^. A woods' road, trail, slashing, overgrown land or an open field all serve in this capacity. The first two are so tenuous a-; to represent increh ;im edge helwecn two * Sec Appendix. |.. 818. A Selecletl by a stntialiralN neeeptable ruiidoMi iiani)>ling nielhod. t See diseuanion on KITerl of Openingv on Nest Location, p. 131!. THE VALUE OF EDGES 173 blocks of forest cover; the rest represent "open" cover types usually of considerable size. These of course also support edge vegetation and may in a sense be looked upon merely as more extensive edges. On the other hand, type boundaries not apt to be characterized by edge vegetation such as those between hardwoods and conifers, appear to influence the location of the nest little if at all. Variation in Importance of Edges The attractiveness of an edge is dependent not alone upon its particular composition, hut also on the makeup of the abutting types. This principle may be stated thus: the attractiveness of an edge is inversely proportional to the extent to which the composition of the types that border it approach the edge in composition characteristics. This idea is aptly illustrated by the nest studies just mentioned. Further evidence to indicate that the richness and variety of the vegetation within a type may exert a strong influence on the use of the surrounding edges by both broods and adults is clearly shown in plotting grouse contacts on the Adirondack study area (figure 15). ALCCn SWAMP HUOWOOOS HARDWOODS a CONIFERS HARDWOODS CUT- OVER (SLASHINGS! FIGURE 15. DISTRIBUTION OF ADULT GROUSE FLUSHES ON THE ADIRONDACK STUDY AREA SEASONS, 1932-1941. each dot represents THREE FLUSHES ALL 174 COVER CHARACTERISTICS AND SHELTER REQilREMEXTS Located in a mountainous region, the soil of the eastern half is sandy and of low fertility. Undergrowth and ground cover are neither widely \aried as to species, nor luxurious as to growth. Slopes are steeper and the larger, mature hardwoods and conifers, which form an often sparse crt)wn cover, attract but few grouse. By contrast, the western half is flatter and more fertile with a fair interspersion of cover types characterized by a much greater abundance and variety of vegetation, particularly undergrowth. A glance at the figure will show that, in the poorer, eastern half, the birds are few in number and are much more likely to be found close to an edge than they are in the western half of the area. The apparent liking of the birds for cover adjacent to roads in the eastern section fits into the general picture. Early settlers laid these out where the grade was easier and where the soil was apt to be better than on the steeper slopes. Here the soil is more fertile, the cover more varied and the birds find conditions more to their liking. The correlation between low- producing cover types and the use of edges by grouse, is clearly evident. One further study, while not in all respects directly applicable here, should be mentioned. It involved the use made by adult grouse and other game of areas reforested with conifers. The Luther Preserve* was selected for this study. This area is composed of some 6500 acres af land reforested with conifers of various species, interspersed with natural woodlands and with open or overgrown fields. The plantations, on the part studied, were arranged in large blocks with the individual pine and spruce planted six feet apart. The average survival of planted trees was about 8.5 per cent. The stand was from 20 to 25 years of age and had never been thinned. The purpose of the study was to determine the use made by game of such coniferous plan- tations in comparison with adjacent natural woodlands. The results as summarized in table 18 indicate that, both in winter and in summer, grouse were much more widely spread throughout the natural woods than in the coniferous plantation. In the latter twice as many grouse were to be found within 200 feet of an edge as were flushed in the entire interior of the plantation. The reverse was the case in the adjacent natural woodlands. TABLE 18. REL.\TION OF GROUSE FLUSHES TO EDGES IN PLANTED CONIFEROUS PLANTATION AND IN NATURAL WOODLANDS- LUTHER PRESERVE— 1933-1934 Percentage of flushes per 100 acres of cover type Id natural woodlands In planted conifers 0-200 feet from lypf iMlge OviT 200 foet from typo edge 0-200 fcot from type edge Over 200 feet from ty[)e edge 34 73 62 S6 66 27 Winter 38 One must remember that the grouse distribution here indicated represents not alone the attractiveness of the edge but also that of the adjacent type. On areas reforested to conifers, edges are of prime importance to grouse since the) must jirovide the varielv of vegetation lacking in the plantations themselves. This situation has not, as yet, been widely grasped eillu-r by foresters or by wildlife managers, who still think largely * In Saraluga County. I THE VALVE OF EDGES 175 in terms of gross coziiposition rather than the mosaic of vegetation which may be produced by cuhural operations. Reviewing the whole, the picture takes shape. \^Tiere the soil and the other site condi- tions are poor, both the variety and interspersion of plants suffer accordingly. Here edges are important in that they add diversity to the cover. Where the soil is good, the land not too dry and the environment has not been heavily impoverished by man or by natural agen- cies such as fire. Nature encourages a wide variety of plant species. In such situations the individual types usually provide within their borders the conditions that strongly attract grouse thereto. Thus there is less inclination to frequent the edges. When fresh desire or seasonal changes draw them to another type, good grouse habitats are usually so well broken up that the birds find no particular reason to stop at the edges. This, probably, explains the pres- ence of many birds scattered throughout, rather than along, the edges of a type on the three study areas characterized bv a favorable and frequent interspersion of cover. ^Air>. ,■• THE T.liTHKR PRE.SERVK IS REFORESTED WITH BLOCKS OF CONIFERS WHICH OFTEN ADJOIN NATURAL WOODLANDS Throughout this discussion we have used the word "edges' advisedly. Plotted on the cover maps are all the major vegetative groups occupying more than a tenth of an acre which are sufficiently distinct to make recognition of edges practical. Within these are found subdi- visions in infinite variety. Here, in a wet spot, a patch of jewelweed has established itself; there, where the winds have tipped over a widespreading beech, letting in a shaft of sunlight, a patch of briers is springing up. Such spots cannot be mapped and measured as one does the cover types. Yet each potentially may fulfill some grouse need. Each has an edge which may be meaningful to a grouse even though too small to map. The more varied the patch- work, the greater the chance for the cover to fulfill the wants of the bird. 170 COf ER CHARACTERISTICS AND SHELTER REQUIREMENTS We have niedtioned size and lompositioii of the edges but not their shape. The late Harry Rogers, dean of game breeders in this country, long ago pointed out that half a dozen rows of corn along a swale would not be as intensively used by pheasants as though the feed patch was 15 to 20 rows wide. A narrow fringe of shrubs between woodland and brushland is. as we have seen, less likely to be used by grouse than are either of the adjacent types. Thus it appears that width may also influence edge use. To such an idea, one can think of exceptions. In classifying these one finds that they are largelv valid only when either one or both of the abutting types are markedly poorer in food and shelter quality tiian is the edge cover. "^ GOOD "edge" cover To sum up, grouse cover productivity seems to he more nearly proportional to the variet) of the composition within each type making up the covert, than to the amount of type peri- |)hery, unless the cover is so poor that the edges furnish markedly greater food and shelter values than are to be found within one or both of the adjacent cover groups. VALUE OF CLEAR-CUT AREAS* In the foregoing discussion of grouse cover requirements during each of the three main life periods of the birds, it lias been shown that cut-over areas or slashings are among the more important types which may enter into the composition of an adequate habitat. Their pri- mary function seems to be that of serving as summer and early fall feeding grounds for broods and adults. They also represent openings which act to break up extensive tracts of woodland. Furthermore, their influence becomes efTective within a sliorl lime after thi\ are created, whereas plantings require many years. Cut-over areas, therefore, oiler the game manager a tool promising relali\cl) quick results • Wrilirn hy Rul.rrt Durrow Jur to the nbicnfe of iho lenior «iillHir in military tprvicr. VALUE OF CLEAR-CUT AREAS 177 where existing woodlands need to be broken uj) or where adequate summer feeding grounds are lacking. To study their role in creating and maintaining cover favorable to grouse, a number were cut on the Connecticut Hill area although nowhere was this area seriously in- adequate in this respect. During the winter of 1932-33 a series of 2-acre blocks were clear-cut in the woodlands 0i 4i ; SMALL SLASHINGS ARE ATTRACTIVE; TO BROODS AND ADL LTS ALIKE on a group of conqjartments adjoining the area. Most of these slashings were located al points over 300 feet from recognized existing openings. The following winter a number of slash lanes, from 30 to 50 feet wide and of varying lengths, were created. All were then left to follow the natural course of forest succession. The climax flora of the Connecticut Hill area is northern hardwood with beech, maple and hemlock predominant. After clear-cutting on such a site the duff of the forest floor dries out. killing most of the shade-tolerant herbs and shallow-rooted seedlings. At the same time the increased amount of sunshine reaching the ground stimulates the stored seed of many species, such as raspberries and cherries. The conditions thus created, especially when the brush has been burned, also enable the wind-borne seeds of such pioneers as aspen and birch to be- come established. Beyond this an abundance of sprouts from the newly-cut stumps soon springs up. As time goes on the tem])orary growth is gradually replaced by the more per- 17f! COVER CHARACTERISTICS AM) SHELTER REQUIREMENTS manent trees and shrubs of the cliiTuix type as outlined in the discussion of succession (p. 118). During the first growing season a winter-cut slashing is little more than a bare opening in the woods. Although a few herbs usually flourish and some stump suckers spring up, the briers, shrubs and seedlings have not yet had time to take hold. By the second summer, however, the new growth has attained an appreciable density affording both food and shel- ter during this season. This density, and the variety of plants as well, increase until about the sixth year and are accompanied by the occurrence of a high abundance of insects, espe- cially in the low undergrowth zone*. After that, development becomes largely a matter of added height and the filling in of the overhead canopy. This results in a gradual shading out of any intolerant low-growing forms. Thus by about the twelfth to fifteenth year the cover, in most cases, consists predominantly of a thick stand of saplings, characterized by sparse undergrowth and little or no ground cover. If not recut it will rapidly pass into sec- ond-growth woodland. Aside from the fact that they created openings in tracts of woodland where such features previously were lacking, these slashings seemed to exert little influence the first year. During the second, and more particularly the third growing seasons, however, they were frequented to a noticeable degree by both broods and adults during the summer and early fall. They remained highly attractive through the eighth or ninth year after cutting, then gradually di- minished. Their use during the spring has been less marked although greater than during the late fall and winter. In spite of the fact that these slashings have proven attractive to many grouse, both broods and adults, their development did not result in any pronounced shift in the distribution of the birds in the compartments involved. True, in many instances broods, in order to visit a slashing, extended their territories through portions of the woodlands where they probably would not otherwise have gone. But they also continued to occupy those portions of their habitat which had been most used previously. In other instances, slashings were apparently placed where the original interspersion of cover types had been satisfactory so that, even though broods were found consistently in their vicinity, records reveal that the same situa- tions had also been occupied in the preceding years. Neither was survival among the chicks of the broods frequenting these cut-over plots consistently better than among those elsewhere on the area. Although this trial did not result in increased productivity one should not conclude that this management technique is not sound. Rather it seems evident that, in general, the diver- sification of the tracts of woodland on this area was suflBcient, without creating additional clear-cut areas, to satisfy the summer territorial requirements of the birds which survived the winter and nested successfully from year to year. There seems little doubt that, on areas supporting low quality cover, the creation of small slashings in the interior of extensive tracts of woodland would result in raising the carrying capacity. In many cases, however, it might also be necessary to improve the winter cover present in order to realize this benefit, since cut- over units of this kind function primarily in fulfilling the birds" summer and earl\ fall re- quirements. On the other hand, such cuttings, although furnishing little winter shelter in themselves, might, if placed within overly-extensive tracts of the latter, enable them to winter a larger number of birds. * See Appeadiz. p. 776. VALUE OF CLEAR-CUT AREAS 179 A further highly important use of slashing operations is to balance, within the covert, the constant tendency for open and overgrown lands to revert to woodland. As has been pointed out, a productive grouse habitat must include openings of one sort or another. Left to themselves, even the, at present, highly productive coverts, occurring where fanning has recently been abandoned, would eventually grow into a continuous woodland of relatively low productivity for grouse. In maintaining the interspersion of cover in such areas neces- sary to encourage grouse abundance clear-cutting is one of the game manager's most useful tools. MMv'T > T^^ ,iK\ /V.i- 'Z'^ ^ AJ'M-- . -- u\, CHAPTER IV FOOD HABITS AND REQUIREMENTS By Gardiner Bump and Jon\ C. Jones STUDIES OF GROUSE FOODS FEEDING HABITS Feeding Periods — Feeding Peciliarities — Caution in Accepting New Foods — Amount of Food Eaten — Variety of Choice — Gravel Consumption — Foods Often Considered Injurious — Effect of Certain Poisonous Chemicals. DIET Foods Eaten and Families Represented — Plant Fouds — Families — rose — birch — wil- low— beech — heath — sumach — sedge — honeysuckle — maple — composite — dogwood — lent — buckwheat — saxifrage — grape — grass — -buttercup — madder — legume — jewel-weed — An- imal Foods — Seasonal Differences in Food Hauits — Regional Variations in Foods Eaten — Yearly Variations in Consumption of Food Items — Variations in Diet Throughout the Range — Food ok the Young — Principal Chick Foods — Monthly Variations in Summer Chick Foods. DISTRIBUTION AND HABITAT REQUIREMENTS OF GROUSE FOOD PLANTS FOOD AVAILABILITY AND SHORTAGES EFFECT OF FOOD ON GROUSE DISTRIBUTION COMPETITION WITH OTHER SPECIES FOR FOOD EFFECT OF OPENING UP THE WOODLAND ON FOOD SUPPLY Natural Openings — Man-made Openings. FOOD AND FLAVOR FOOD AND ITS RELATION TO HEALTH Chemical Composition of Some Grouse Foods — Crude Protein — Fats — Nitrogen-free Extract — Crude Fiber — Ash — Vitamins — ^Differences in Chemical Composition — Re- lation ok Food to Weight — Relation ok Food to Cycles. WATER REQUIREMENTS 182 FOOD HABITS AND REQUIREMENTS SUMMARY Food shortages severe enough to cause star\'ation. pronounced weight losses and death apparently do not occur, (p. 229). The ruffed grouse is omnivorous, (p. 184). It is primarily a plant-eater, its known consumption including representatives of 65 fami- lies and over 334 species, (p. 197, 850). Manv of these species are widely distributed: they are normally to be found, often in abundance, in good grouse coverts, (p. 1981. For the most part these are pioneer and temporary (transition) type species, (p. 199). Trees and shrubs furnish the bulk of the food. (p. 198). Few of the commonly eaten food species are found in dense woodlands, (p. 225, 231). The maximum number of these plants is present three to five years after heavy lum- bering, (p. 234). Animal food, mainlv insects, is predominantly taken only during the first two weeks after hatching, (p. 212). With adults animal food makes up but one-twenty-fifth of the summer diet: during other seasons it is much less. (p. 197, 212) . Available insects during the critical period following hatching may reach 330,000 per acre in areas recently lumbered, (p. 214. 776). Insects most frequently taken are ants, beetles and caterpillars, (p. 213). Over an 11-year period the foods most commonly taken by New \ork grouse came from aspens, cherries, birches, raspberries and blackberries, hop-hornbeams and thorn- apples, (p. 215). The diet varies year by year and season by season. Some of these variatioiir^. but not all, may be explained on the basis of the relative abundance of the foods, (p. 214, 218). Great differences in diet exist between various parts of the grouse range. The young differ widely from their parents in food eaten at first but acquire adult tastes by the end of August, (p. 220, 225). To obtain an accurate conception of food consumption over a large area, such as New York, a minimum of 50 grouse should be examined each season during each of at least five years. However, a fairly large list of foods taken can be compiled from the results of a single year's study. (p. 184). The distribution of grouse is influenced to no small extent by the feeding grounds in summer and fall. (p. 229). Normally grouse find satisfactory amounts of water regardless of the specific habitat they oc- cupy, (p. 245). Sfc^ STUDIES OF GROUSE FOODS 183 Food is one of the cardinal factors that determine the abundance of any wildlife species. Food shortages have long been considered a paramount cause of wildlife scar- city. Corrective measures, for the most part, have been quite inadequate, although usual- ly dramatic enough to appeal to nature lover and sportsman alike. As a result, studies of grouse food habits, ranging all the way from the penknife ex- amination of a bird killed in the field to the most careful laboratory analysis, have furn- ished an informative but haphazard picture of what grouse eat at various periods and over many parts of their range. STUDIES OF GROUSE FOODS The first extensive survey of grouse food habits was Sylvester D. Judd's analysis of 208 crops and stomachs gathered from widely separated parts of the bird's range. Fall food habits of New England grouse have been admirably described by Gross' '^ Nelson, Clarke and Bailey""" have published the results of a small but thorough sampling of adults collect- ed in Novemlier and December in the George Washington National Forest, largely in Vir- ginia. Under the direction of Dr. A. A. Allen. Thomas Smyth"" prepared an excellent thesis on the food habits of the ruffed grouse I mostly fall and winter adults I with special emphasis on the region about Ithaca. N. Y. Local reports have been produced by MacGregor"" for New Hampshire. Kuhn"'" for Pennsylvania, and Chaddock"' for Wisconsin. Unfortunately most investigators have been largely dependent for specimens on birds shot during the hunting season; thus there is a dearth of infurinalidii fnr oilier periods of the year. The food of grouse chicks has received little attention ex(e]il li\ llic i iirrenl study. The situation nia\ best be visualized by glancing ihrmigli table 19 identifying the more comprehensive grouse food investigations. TABLE 19. MAJOR STUDIES OF THE FOOD HABITS OF RUFFED GROUSE Inv('stiKJitf>r Nutnhcr fxtitniricd Ap:o of birds .Si'nsons studied Collection area Years covered Datji on occurrence Data on volume Judd, S. D 208 R Adults Chicks Largely the colder part of the year. June Mostly the North- east. Not ffiven X Smylli, T Ml II Adults Chicks Mostly f.ill iiii.l winter. Chiellv from vicinity of IlhacM. N. V. Mostly 1922-2.1 X X Gross, A. 0 1 ,o.';o Adults Oct -lice. New ICric. & Mari- time Provinces of Catuida. 1926-32 X Nelson, A. L.. Clnrke, T. E., and Hailoy, W. W 18.''> Adults Nov. -Dec. Ceo. Wash. Nat. Forest, Va. & W. Va. 19.').i-.16 X X Kuhn, T. M 2.!0 Adults Fall. Pa. Mostly 19,i8 X X Hninp. (i.. iirid Jon(\s, J. C 1,(10,'! Adults Chicks 12 months. Juric-Aiij:. New York New York 1931-41 1931-41 X X s^ Kelso, I,. H. 80* 111 Adults Adults Dec-Mar. Dec- Mar. New York Northeast 1931-31 X X Miu;(tregor, A. K., et al 1(12 2:16 Adults & Chicks Adults June-Nov. Mostly Sept., Oct. & Nov. So. N. H. Wisconsin 1910 19.38-39 X X X Chadiloek, T. T X PcldiTlioi-r. E. B 176 A Unknown 12 months Iowa 1938-40 X •Collected by the N. Y. S. Ruffed Grouse Investigation and examined for it by the U. S. Fish and Wildlife Service. They are accordingly utilized in compiling the results of the Investigation's food habits work. ADroppings only examined. 184 FOOD HABITS AND REQUIREMENTS Adequate support has allowed the Investifration to look into the food habits of grouse in New York on a comprehensive scale. Experience indicates that these habits may change materially from year to year and from season to season. The habitat from which a bird is collected, as well as its age (whether adult or chick) may often be surmised from the character of the food. Another factor is the variation in the distribution and abundance of grouse foods, not only as between such extremes of range as Alaska and Alabama, but also to a surprisingly marked degree within the single State of New York. To determine the differences between food habits for regions and habitats, years and seasons, adults and chicks, it has thus been necessary to collect specimens over a period of 12 years and from three more or less differentiated regions of the State. To indicate reli- ably changes in the food taken, a goal was set for each year of at least 75 adult birds to be examined from each region during each of the four seasons. Similarly, at least 50 chicks were collected representing each region for each of the months of June, July and August. (See table 20). TABLE 20. SKASONAL AND RKGIONM. DISTRIBUTION OF THE l.OQ.T ADUIT GROUSE AND 510 CHICKS, COLLECTED AND ANALYZED FOB FOOD HABITS STUDIES— ENTIRE STATE— 1931-1941 Region Adults Chicks Season Month Spring Summer Fall Winter Total June July August Total A an excusable fallacy to consider llial food research may well stop when the FEEDING HABITS 185 details of stomach analyses are in hand. This impression is heightened when one finds that food shortages apparently are not directly limiting influences on populations of the rufFed grouse. In fact, the Investigation has yet to examine its first bird, the death of which could be attributed to starvation. Upon deeper analysis, however, less apparent food rela- tionships are revealed which, in the long run, may affect grouse welfare much more than does an occasional empty stomach. As with wildlife in general, even the simpler inter- relationships of food with shelter, with predators, with diseases, with cycles and with the ability of the species to reproduce, have heretofore been little recognized nor much investi- gated. Not enough thought has been given to these less familiar effects in connection with the study of food habits and requirements. Hence the Investigation has not limited its work to a determination of what grouse eat nor of how much of each food item is consumed. Some of the correlations referred to are so complex and little understood as to baffle understanding. That the composition of sunlight may differ from year to year is a recog- nized fact. But to j)rove, with ])reseiit knowledge and equipment, that such variation is, or is not, a basic cause of periodic changes in abundance, as suggested by DeLury"" and King, is an undertaking, challenging indeed, but probably beyond today's scientific re- sources. A start, however, has been made by measuring the chemical composition of some of the more commonly eaten grouse foods. These, together with |)i'itincnt data alreadv pub- lished, are presented later.* Some attem|)t has also been made to determine (he nutritive requirements of grouse in captivity. Thus in 19.'?-!. and 19.3.5 Dr. L. C. Norris. working for the Iiivcsligation. found that young grouse thrived best on a starting diet of 27-.'?fl per cent protein wliich cnuld be safely reduced to a 20 ])er cent level at eight weeks. For (he first time light has been cast on the extent to which common grouse foods satisfy protein and other needs by a study of their effects on tlic metabolic rate of captive birds. In this way some idea of the nutritive value of the more important foods has been gained. Empirical knowledge of this sort, while basic, must be translated into action in order to realize its potential value. Thus, when the txpical diet became known, the next step was to chart the distribution and approximate abundance of these foods throughout the State and the habitats in which they most commonly occur. Nature is a generous provider. Man has partially learned the trick of using her bounty for his ends. Thus some time was spent in learning how to produce the most desirable foods in quantity and to make them grow where they will do the most good in creating more productive grouse habitat. FEEDING HABITS'^' One cannot long watch either an adult grouse feeding or a brood fanned out in advance of their mother and on the alert for food without sensing that these birds have absorbed something of the dignity and quiet of the woods in which thev live. While quick and wide- awake, the birds show little of the nervous activity so characteristic of the bobwhite. Feed- ing is for the most part a leisurely pastime, for food is plentiful and seldom difficult to obtain. * See disciissinn of Clieinical Conijiosition of Some Crousr Foods, p. 237. A The anions of yroiisr. both ji. lulls and >otiiig. wliilt- f.-ciliiie. arc described in Chapter V. 186 FOOD HABITS AM) REQUIREMENTS Feeding Periods Many novices at hunting grouse are sure that the birds feed at the beginning and end of the day. Oldtimers are more reticent about committing themselves. Each can likely re- member seeing a cluster of birds about a thornapple tree in late afternoon, or birds budding an apple tree at woods edge well into the evening when the moon was bright. But the oldtimer also remembers grouse feeding at high noon on beechnuts shucked out over the snow on a January day. The truth seems to be that where particularly attractive foods are to be found, there is a tendency for the birds to feed early and late: but the exceptions are legion. In general there is so much food available in most New York grouse habitats during all seasons of the year that the birds can feed al any time they choose. Where favorite foods abound the birds must occasionally spend hours filling their crops with unbelievable amounts of buds, leaves, catkins or fruits. Thus one bird collected in win- ter was found to have eaten 1.300 buds of trembling aspen (Populus tremuloidesj , while another had taken some 1.400 buds of cherry (Pruiius) as well as smaller amounts of three other kinds of buds. Since hop-hornbeam (Oslrya virginiana ) catkins are by no means as common as the buds, the grouse that ate 820 of them must have spent a substantial portion of a day or night aloft in the quest. The same may be said of the bird in whose crop 1.069 hairy fruits of staghorn sumach (Rhus typhina) were found. Nor is this filling-up tendency limited to winter, for the crop of a bird collected in spring in the Catskills contained nearly a half pint of mountain laurel (Kalmia lalijolia) leaves. A crop obtained in the same region in summer held 20 fruits and more than 6.000 seeds of strawberry ( Fragaria). A favorite trick of both chicks and adults is to combine the lazy luxury of a summer dust bath with the picking uj) of insects, [larticularly ants, that may venture their way. In summer, though, food seems to be so abundant that the birds can get all thev want at any time, for the crops are not so well filled, on the average, as at other seasons. (See table 211. TABLE 21. AVI:H\(;I-: \()1J \II;s, I.\ cubic CKNTlMI-n'KHS, OK Till-; CROP \M) C.IZZABD CONTENTS OK 1.0'):i ADULT CHOUSE, BY SEASON AND REGION— NEW YORK Spring Summer Fall Winter Crop Ciizzard Crop Gizzard Crop Gizzard Crop Gizzard 17.5 24.7 18.0 9.8 10..1 8.9 5.0 .1.5 6.1 6.5 5.9 7.7 15..3 15.9 10.5 7.5 8.1 6.9 .16.9 21.0 16.2 10.9 CnUkillH 9.7 Rest of State 9.2 20.1 9.6 4.9 6.7 13.9 7.4 24.7 9.8 'Average for all years, all seasons: crop. 15.9; gizzard, 8.4. Of all the elements, onlv wind seems coniinorii\ In ;i|> with whole leaves of poplar or white oak acorns. Chestnuts, in the old davs before the l)light. were a popular food item in the fall, as are still the whole fruits of th<)rMa|)plc. riicsc arc dftcti tlic width of a good-sized thumbnail. Stories of grouse swalldwing \oung mice have been received but not verified hv the Investigation, though I'olderboer after cxaniitiiiig Iowa grouse droppings, reported one or two collected in summer to conlaiii bones ami hair of deer mice ( Pfroiuyscus). Judd'^ found a green snake 17 inches long in one grouse crop, indicating that the birds are deterred h\ neither shape nor length once an item has been marked as food. In computing the bulk contents of the cro]i> and gizzards by seasons, as indicated in table 21, a curious situation was brought to light. There seems to be little significant difference in the amount of food found either in the cro|) or in the gizzard of grouse during the spring, fall or winter months. In summer, however, it is so much lower as to indicate a marked re- duction in food consumption and in the time spent in feeding. The greater digestibility of summer foods, such as soft-bodied insects and flcshv fruits is another contributing factor. C.\LTioN IN Accepting New Food.s With prccocial birds like grouse which leave the nest within a few hours after hatching, the task of di'termitiing what is edible seems to fall largel\ upon the voung. In captivitv at least, the mother, unlike the barnyard hen, seems to spend relatively little time and effort FEEDING HABITS 189 in teaching them what to eat. Perhaps Nature has accordingly provided the chicks with the "pick-up" habit. It would seem all grouse exhibit it more or less throughout life. Chicks seem at first to pick at items which move or are small and contrast with their surroundings. Whether they have a "feel" for those which have food possibilities, whether the mother, in a way too subtle for detection by human eyes or ears, instructs the chicks as to which items, among those picked up, are to be eaten, or whether they learn purely by trial and error, can only be guessed. Certain it is that the chicks are constantly picking up small articles, only a fraction of which are actually eaten. By the time grouse are half grown this early caution in accepting foods has developed into a habit. An instance comes readily to mind. In trapping grouse, a comparatively simple pro- cedure in the deep woods where they are tame, it was found that birds come readily to feed- ing stations established before the ground becomes snowbound. However, once snow forced them to seek their food largely among the treetops, many grouse exhibited a surprising un- willingness to return to a diet of fern leaves, thorna])ple fruits and other presumed delicacies previously eaten. Likewise illustrative is the difficulty experienced in getting captive birds, long used to eating grain and pelleted mash, to take foods that are staple for their wild brethren. Hand-raised grouse may starve rather than change to a diet of the buds so ap- petizing to free-living birds. Five bushels of beechnuts contributed to the Investigation re- mained largely untouched because of the difficulty of convincing some 200 grouse that they were good to eat. There is something deeper than mere caution involved here, however, for the birds. Imth in captivity and at feeding stations, iiurease markedh their consuni|)tion of corn as the weather becomes colder. iNormally little of this grain is taken, but as the temperature ap- proaches zero this tendency is reversed. Here is a problem for the bird physiologist ( and psychologist) to answer. Amount of Food Eaten The daily and seasonal consumption of food by wild grouse is not susceptible of accurate measurement, but conclusions from the record. |)ie(emeal though it is, are not without in- terest and probable value. The daily intake varies with the availability and character of the food, the age of the bird, the weather, and a host of other factors. A chick will take unlit'lic\al>!e amounts of easily digested insects and succulent vegetation in the course of a single day"s feeding. In captivity it requires from six to ten mealworms each an inch long to satisfy a week-old bird. These may be fed at half hour intervals. Two 7-week-old birds raised under semi-wild con- ditions, each proved equal to consuming 30 blueberries, several raspberries and from six to ten medium-sized grasshoppers at a meal. Leaves and buds in surprising quantities were also much relished. The same birds three weeks later were each consuming a head of lettuce as large as themselves each day. The astonishing total of 4,350 strawberry seeds, perhaps rep- resenting 50 fruits, was found in the crop and gizzard of one gluttonous young bird col- lected in August in the Adirondacks. More than one day's food intake, however, was prob- ably here represented. From an August adult in the same region were taken 20 whole strawberries and more than 6,000 seeds. The largest crop examined, containing 153 cc, was that of a bird shot in spring in the Catskills. Nearlv two-thirds of the contents consisted of leaves of mountain laurel (KaJmia latifolia), though there were 288 buds of aspen (Popiihis tremuloides) also present as well 190 FOOD HABITS AND REQUIREMENTS as eight other items in smaller amounts. Together, this crop's contents would constitute a good-sized handful. It is not unusual Im Imd l.irds stuffing themselves on relished foods at any season of the year save summer. Leaves are fa\orites during spring, fruits in early fall and buds and catkins in winter. It was at first surprising, but on second thought logical, to find no cro])s containing more than 19 cc. of food in summer. The diet then is mainly insects, juicy berries and tender THE AMOUNT AND VARIETY OF FOOD FOUND IN THE CROP OF A GROUSE IS OFTEN SURl'RISINO PICTURED IS THE CONTENTS OF ONE WHICH CONTAINED: /. ]HH buds, hop-hornbeam 4. 143 buds, hawthorne 7. one hud, scarlet oak 10. 223 fruits, staphorn siiiikii h I'.i. 37 fruits, seeds, bittersweet I(>. leaves, speedwell 2. U)() catkins, hophornbcani 3. 09b buds, scrviceberry .5. 38 buds, witch hazel 6. 2 buds, red cherry fi. •W buds, sugar niaple 0. 86 buds, low-bush blueberry //. 3 seeds, hawthorne /:'. 4 fruits, seeds, maple-leaped //. leat'es. hawlncffd viburnum 17. leaves, strauberry 15, leaf, common speedwell leaves which digest rapidly. In tin' midst of pltnt\. it is also possible that the birds may feed more fn-ipicnlK and lake less at a time. In coiilrast to ihc crop, the giz/.ard holds a nuirh smaller amount of food. The largest ob- served to date was thai of a winter bird from the ('.atskiil>. ll coiilaiiicd 20 cc. FEEDING HABITS 191 Variety of Choice Perhaps the "pick-up" habit has something to do with the bewildering variety one finds in analyzing grouse crops and gizzards. Certainly this species is one of our most omnivo- rous game birds, for no fewer than 580 animal and 414 plant items have been identified from New York State grouse alone. From one summer bird, 45 separate items were determined. Although the average bird feeds upon a more varied diet during the summer than at any other time of year, there seems to be no period in which a considerable choice is not avail- able. Even in the teeth of a hard Adirondack winter, one grouse found 16 different items. However, the average for 90 birds collected in this region in winter over a period of years was only 4.6 items each. The seasonal and regional variation in number of food items is indicated in table 22. TABLK 22. SEASONAL VARIATION IN THE NUMBER OF FOOD ITEMS PER INDIVIDUAL CROP AND GIZZARD TAKEN BY GROUSE IN NEW YORK AdulU Season Rnf^ioil Spring Summer Fall Winter Maximum Average Maximum Average Maximum Average Maximum Average 20 26 26 5.2 7.0 6.6 6..) 24 .11 45 10.2 11.0 12.8 11.3 17 24 18 7.5 9.5 8.0 8.3 16 Cotakills. 4.6 Rest of State. . 6.6 Average 6.5 5.9 Chicks Month lU^Kiiiii June July August Maximum Average Maximum Average Maximum Average tl 37 U 12.4 17,1 1 II ll.fi 29 :u. II 13.1 Ift.O l,-..o 17 Catskills 9.5 Average 10.4 — _ In spring Nature sets a more varied table for the chicks. One active youngster in June, succeeded in catching 34 different insects and feeding on seven different plants. Fewer in- sects are taken in July and by August the menu of the young approximates that of the brood leader, the mother, in lessened variety if not as yet in increased bulk. As the observant hunter knows, grouse take a toll of many things in its environment. Buds, twigs, fruits, berries, seeds, grains (particularly buckwheat), flowers, leaves, stems and even roots, are all eaten as fancy dictates. It is proper to call these birds budders. browsers, berry-pickers, seed eaters, leaf-clippers and even grain-gleaners, and they may be all six on many a feeding da> . Nor do they find a smaller array of food items among the animals, for eggs, larvae, cocoons, pupae, nymphs and adult insects, as well as insect allies, such as spiders, all are fair game to the hungry grouse. Under stress of circumstances, some unusual items may be taken. There are at least five 192 FOOD HABITS AND REQUIREMENTS published rwords of the birds eating snakes. The 17-inch green snake mentioned by Judd is an example. A slightly smaller garter snake was found in a grouse crop b\ the Investigation. A nunilxT of scales from the same species were likewise identified in two other birds. Four other vertebrates also have been recorded. Bones from a small unidentified bird were found in a grouse shot May 23, 1933 in the Adirondacks and feather remains in that of a Catskill bird. Even more mystifying were the cottontail hairs found in two other gizzards. It is more easily explained but equally interesting that another individual had fed to a slight extent upon small finger-nail clams ( Sphaerium ), and that twice mussels have been recog- nized. These mollusks are more or less common along the marshy shorelines of certain Adi- rondack lakes. Snails also were taken by 75 birds, mostly chicks. There are a few plants in which the grouse exhibits a little-suspected interest. The leaves of pondweed ( Potainogeton), stalks of sedge (Cyperus), seeds of spikerush ( Eleocharis), bits of moss and traces of fungi and of mushrooms, have all been identified once or twice. There are in addition miscellaneous items among which might be iiu-iitioncd: long-dead leaves, birch curls, bits of glass and an inch-long roofing nail (the latter |)uncturing the giz- zard and killing the bird). Lead shot seems rarely to be picked U]). though one case of what appeared to be lead poisoning was investigated. Gravel Consumption Seed-eating birds in general pick up bits of gravel and other hard particles to aid in grinding their food in their unusually strong muscular stomach called the gizzard. It is the function of this organ to pulverize the food before passing it on for digestion in the intestines. As a mechanical aid in this process the grouse, like its near relatives, usually keeps a small supply of gravel or other hard objects in its gizzard. The number of these gizzard stones in a grouse is usually greater than one would suspect. In iNew York birds, they are present in amounts varying from a trace to nearly one-half the total bulk, the average being about 12 per cent, or one-ninth of the contents. It is possible that gravel is necessary to efficient gizzard action, but its absence for short periods at least seems to make little difference. With buds, twigs and hea\ y-coated seeds the breaking down action may be quite successful even in the absence of gravel. In such cases it is not un- common to find a number of hard seeds which, it has been suggested, may act in somewhat the same capacity as grit. If such action does take place, it is of short duration in the ruffed grouse, for seeds in the gizzard seldom show much evidence of long-continued wear. Pebbles, on the other hand, even the resistant granite. riia\ be retained until they are smooth as though hand-polished. The presence of grit in the gizzard, bulli in ficciiicncN of occurrence and in amount, seems to be inlhjcnced more or less by the weather. b\ llic lialiitiil ;iiul b\ the fond which the grouse happens to be eating. Most birds want grit at all seasons of the year. This In noticeable al winlcr feeding stations, both for grouse and for pheasants where grit, as well as food, is taken. Whenever winter blankets the coverts with snow. gr;i\fl is hard to obtain and grouse ma\ retain the fall-secured sup|)l\. This tna\ explain wh\ gravel makes up utcd the ill-effects to failure to draw the birds immediately, and stated that after taking a lumdful of laurel from a '"pheas- ant " he had eaten freely of the meat without any unpleasant consequences. (Arouse were once banned from the markets of Philadelphia through fear of laurel poisoning. Mention has already been made of one grouse consuming a half-pint of mountain laurel leaves at one feeding without any indication of discomfort. Among the birds examined, such leaves were eaten by 23, comprising over half the contents in four. Twigs and buds were present in 11, but usually in small quantities. In an effort to cast additional light on the subject, six adult hand-raised grouse were pro- vided with their normal daily ration supplemented with a continuous supply of fresh laurel leaves. At the end of one week four were autopsied without finding any indication of in- jury that could be ascribed to laurel. The two remaining liirds were continued on the same schedule for three more weeks without visible harmful results. They were then put on a diet of pure laurel. Death resulted. Upon autopsy, it was attributed directly to malnutrition. Two additional grouse were fed on laurel only. Both showed consistent weight loss and died after seven days. Death from malnutrition again was indicated. Remains from six of the grouse, five from the first lot and one from the second, were fed to mice and kittens, none of which showed indications of discomfiture. The other two were eaten by the senior author, who reported no ill effects, though the taste of the flesh was dis- tinctive. Though these tests are admittedly not extensive, one finds here no indication that laurel is poisonous to grouse, or through them to humans. When eaten alone it apparently has little sustaining value. Grouse will eat large quantities of laurel leaves when deprived of other food and freely su|)|)lemcnt their diet with laurel even when pro\ ided with an abun- dance of their normal rations. That the grouse like many other birds can eat the berries of poison ivy (Rhus toxicoden- dron) with ap|)arent inqiunitv is another interesting fact. Sm\th " quotes Forbes as finding 280 of these berries in a bird collected in December in Illinois. Dr. A. K. Fisher counted 160 which were in the crop of an Adirondack grouse shot in October. .Nine of the adults and one chick collected for the present Investigation had been feeding on jioison ivy fruits or seeds. That even ii] the stomach they may retain jiroperties poisonous to man was indicated by Judd. who cited the case of an investigator who was poisoned uliilc cxainiiiing the gizzards of crows that had fed on poison ivy berries. A number of other plants contain substances poisonous or highlv unpaialalijc to man. Some of these are useful grouse food plants, while others are conspicuous In their absence in the normal diet. The berries of yew (Taxus canadensis) and fruits of bittersweet (Celaslrus scfindrns) contain toxic alkaloids, but that did not deter 20 birds from eating the former and h\c llic lallcr. Seeds of the smartweeds (Polygonum ) ina\ cause |>oisoning when fed in grain or ground fc('\e experiment was repeated using a standard solution of two level teaspoonfuls to one gallon of water. None of these birds showed ill effects. Grouse were also found to be surprisingly resistant to powdered strychnine. The minimum lethal dose of most humans is 0.5 of a milligram per kilogram* of body weight. Three birds were fed respectively 92. 108 and 126 milligrams per kilogram of body weight. All survived. The amount of strychnine consumed was respectively equal to 184. 216 and 2.52 times the lethal dose for humans. Although the birds were found to be tolerant of powdered strychnine. 80 milligrams of the same chemical in solution caused death within a few minutes. .Strychnine does not mix readily with water so that solutions such as here tested are seldom used. DIET In considering the food of grouse it is common to ascribe too much importance to the species most commonly eaten. It is generally believed that the birds exercise a choice of food items based on need. Were this the storv. food habits research would be tremcndouslv simplified, for one would have onl\ to determine the relative consumption of various foods by seasons, regions and \cars for adults and chicks. Knuwing the volume of each food eaten, one could then easilv cinnpilc a list of plant and aninial species to be encouraged. It is not as simple as that, for tiicrc are rea-nlitx) ( '.hcrrv ( Pniiiii.i) Krrh'i lift Ilia) BlaiklMTpy and rMS[)l>eTry (Rtibus). . Hop-hornlieiiiii (O.slrvd) Thornapplc {( 'niliii'dus) Strnwl)('rry (Frwiaria) Apple (A/n/u,«) Beech (Fmius) Sumach ( Hhiis) Sliii(ll)iisli ( \ inelanchier) Scdfje (Carex) Vihurmim (Viburnum) Maple (Acer) Do;;wo(»l (Cornus) ( )ak lOucrcus) Laurel ( l\iihiii 2.3 2.0 2.0 \.h l.ft 1.1 1.2 1.0 1.0 1.0 .<) .K .5 IrnceA .1 .1 Number of birds ID which found* 2lt 118 232 282 112 :!i I l'»T 81 101 2.'>l 101 2(1.3 I 12 I7>» ()1 30 63 75 87 88 16 26 16 213 312 226 43 79 71 68 123 13 •Arliiiil niiiiihrr (jf occurmirrs of (he k<'i>iik or onlrr; niniiy binU hud entoii murf Ihnii nun stprrirs. ALciUi tliuii 0.1 |HT cent. DIET 199 These families contain many of the pioneer species which fill in old meadows, pastures, or slashings. Several of the species contributing to grouse food as the blackberry, cherry, sum- ach, viburnum and aspen, are among the most common and widespread of northeastern woody plants. Within each family are from one to a dozen species which, under the right condi- tions of time and place, may furnish large quantities of buds, leaves, fruits or seeds on which many of the grouse examined have fed. Let us then look into these families a bit. The Rose Family. One of the most common and important of these families is the rose group (Rosaceae). During warm weather, strawberries, raspberries, sedges and aspen are the most commonly sought after food species. Of these, the first two belong to this family. In the late spring and early summer, leaves and fruits of strawberry (Fragaria) are prime foods for both young and adults in all parts of the State. A chick collected in the Southern Tier in July contained 1,650 seeds; in another from the Adirondacks. shot during the same month, were found 2,100 seeds. Adults likewise relish strawberries, as is evidenced by a Catskill bird which had eaten fruits sufficient to provide the more than 6,000 seeds that were found in its crop and gizzard. By the time the strawberry moon wanes, the raspberry and blackberry iRuhus) season is approaching and the birds take full advantage of it. These plants are also members of the family Rosaceae. It is not uncommon U> find in excess of 2,000 seeds, representing some 30 fruits in a summer crop (and gizzard) of either chick or adult. One youngster collected in July in the Catskills contained .3.160 seeds. What their appetites may bo when they grow up is indicated by the meal of a summer bird from Tompkins County in which more than 8,000 seeds (120 fruits) in the gizzard and 2.'i fruits in the cro|) were counted. In captivity. 108 red raspberries were consumed within half an hour by one individual less than three months old. Occasionally dried fruits, as well as leaves and buds, arc taken throughout the fall and winter months until April. Products of cherries (Primus), probably arc the most consistently sought of all those of the rose family. The hard seeds are picked up throughout the year. As soon as the light red fruits of ])in cherry and the darker drupes of choke and of wild black cherries become available the grouse seeks them out; in one October bird 180 of the first-named were found. In September the wine-red juice of the wild black cherry may impart its color to the entire crop contents. Later the buds are very attractive. 1.400 being recorded from a February bird collected in St. Lawrence County (N.Y.). The leaves, however, are entirely neglected. Although not so widespread in grouse covers, as some of the native plants, the apple (Malus) is utilized the year around. Young birds raised in captivity at the Research Center seemed to prefer ai)|)le leaves to any others offered tliciu. According to sloinach analyses. 200 FOOD HABITS AND REQUIREMENTS these leaves are not much taken in the wild though one bird collected from Jefferson County in late October had fed exclusively upon them. They stay green in the face of fall frosts and this may attract grouse to the trees though they feed largely on the buds and decaying fruit. That heavy pruning of apple buds may result is indicated by the meal of a Catskill bird which had eaten 566 of them in November. In the late summer and fall, the birds have a tendency to gorge themselves on thornapple (Crataegus) fruits. From the crop of one full fed grouse, collected October 16. near Ithaca (N.Y.), were extracted 125 of these bulky fruits and seeds along with 11 other kinds of food. In fact, throughout the State more fall birds were found to have fed exclusively on redhaws than on anv other item. Buds of thornapples, however, do not seem to be much relished and only three birds had eaten the leaves. Shadbush ( Amelanchier ) is also much patronized both for its early fruits and for buds. One winter-bound bird in the Adirondacks gathered 846 of the latter and another in the Cat- skills picked more than 1,300. One of the largest crops examined was filled with almost half a pint of leaves of the barren strawberry (Waldsteinia) ; this bird was collected on December 2. Though avens (Geum) seeds, also of the rose group, were not taken in sufficient quantities to rank among the more common summer and fall foods, one bird taken October 22 in Albany County made most of its meal of them, taking more than 2.000. Occasionally some species with apparently desirable fruits are neglected. This seems to be the case with the chokeberries (Aronia). the buds of which were eaten by only 12 of the grouse examined, seeds and fruits by three adults and one chick: and leaves by one adult. With one exrejjtion, all of these were represented by small amounts. The Birch Fainily. Second in importance, though little patronized in summer, is the birch family (Betulaceae). Chief among the group are the birches (Betula) themselves, among which the birds are so frequently found as to give rise to the appelation of birch partridge. The seeds, though common travelers over the March snow, are seldom touched, but both catkins and buds are well liked. These are taken from all species; those of yellow birch ( B. Ititea). ■v^^^^r:- DIET 201 black birch (B. tenia), and paper birch (B. papyrijera) in largest quantities. Two birds collected from the Adirondacks, one in winter, the other in early spring, had consumed 728 and 1.150 buds and twig tips respectively. Even in the presence of an abundance of other appetizing foods, birch buds are taken as early as September 18 and as late as May 11. To a forester, the hornbeams or ironwoods are weed trees, but every grouse knows they have a place in its favorite covert. The leaves of ths hop-hornbeam or hardback (Ostrya virgini- ana) are but seldom eaten and the early maturing fruits are utilized less than might be expected. The smooth brown buds and particularly the flat-clustered staminate catkins furn- ish a wealth of food which bulks large in many a grouse crop from October through April. Up to 523 of the former and 820 of the latter have been identified from separate birds. The smaller blue beech or hornbeam (Carpinus caroliniana) is browsed less commonly, perhaps because of its more slender branches. One shrub, the hazelnut (Corylus). also a member of the birch family, is sometimes visited by grouse in search of buds or of the staminate anieiits borne at the end of last year's twigs. More adventurous birds may even assay to swallow the large slightly-flattened fruits. The Willow Family. Among the trees and shrubs no family is more widely distributed than are the willows and poplars (Salicaceae). It is, then, fortunate that grouse browse upon them. The large buds, long catkins, and tough leaves set on the stout twigs of the aspens (Populus) are [jarticular favorites even when other food is abundant. Bendire" notes, "The Canadian ruffed grouse, according to Mr. Hardy, feeds not alone on the ])oplar buds but also on the hard old leaves. He writes me, 'I have killed one with his croji filled with such leaves on the 20th of August, and they eat them continuously until the leaves have fallen in late October.' " Green leaves, too. are not overlooked, for they had been commonly taken by the birds studied from April 21 until late November. Leaf-browsing thus overlaps the bud- ding period at both ends. In the Adirondacks, aspen budding begins as early as September 18 and continues at least until May 17. On January 6. one bird made a full meal of 1,300 trembling aspen (P. tremuloides) buds and twigs. Its larger-leaved cousin (P. grandiden- lata) is equally attractive. The willows (Salix). though seldom important as a source of food are not entirely passed by. Of this group, grouse, in common with children, may prefer the pussy willow (Salix discolor), judging from the 636 buds picked off by one bird collected on December 13. The Beech Family. Were it not that abundant crops of beechnuts and acorns are produced only once in three or four years, the beech family (Fagaceae) would be outstanding as a producer of fall and winter grouse food (table 26). In the Northeast, coincident with the falling of the leaves, grouse-wise hunters often head for the beech ridges if it be a mast year. There the birds may have been feeding among the beeches (Fagiis grandijolia) since early September. For the next two months, beechnuts, close-housed in their burrs or shucked out on the snow, are seldom overlooked. But it is the previous fall's crop revealed by March and April's shrinking snows that are really appreciated. Following a beechnut year, they rank sixth in the list of spring foods. Occasionally these nuts are eaten in numbers. Bendire'" mentions "finding 76 in one bird's crop and 60 in another". These were Canadian birds. Beech is a common tree in New York but no such heavy utilization has been found by the Investigation, though Smyth*" records finding one crop collected December 8, that contained 78 nuts. While occasionally found as the sole cro]) contents, beechnuts are commonlv eaten along with other items. One bird col- lected near Elniira on November 5 had gathered 20 fruits of highbush cranberry (Viburnum 202 FOOD HABITS A\D REQIIREMENTS opulus), 42 fruits of arrowwood (f'irburnum dentatum) and ten other items, including 25 beechnuts. TABLK 26. QUANTITIES OF BEECHNUTS CONSUMED BY 1,093 ADUl.T GROUSE IN NEW YORK— 1931-1941 Occurrence of beecbnute Nunib<»r of birds Year examined Number of birds Percent of Volumetric in which found occurrence percentage 1931 23 1 4.4 0.1 1932 30 •> 6.7 0.7 1933 94 4 4.3 0.4 1934 164 36 21.3 8.3 1933 302 31 10.3 1.9 1936 141 1 0.7 0.2 1937 78 11 14.1 8.5 1938 19 0 0.0 0.0 1939 88 4 4.3 3.2 1940 97 9 9.3 2.5 1941 57 2 3.5 1.0 In years of beech mast scarcity grouse of course turn to other foods. The long spiked beech buds, too. are occasionally eaten, though seldom in quantities and then probably in- cidental to feeding on the nuts, for they are usually more common in early spring and fall crops. One bird collected in October in the Catskills. however, contained 601 of these stick- like buds. Beech leaves also are taken occasionally. Now that the chestnut has all but vanished, the other great group of mast producers upon which wildlife must depend is the oaks (Quercus). Not such a factor in grouse economy in the Northeast as are beechnuts, acorns may nevertheless be taken in sizeable quantities. One bird killed October 15 had eaten ten of these large nuts. Acorns from the white oak group, particularly those from the white oak iQ. alba) and the bur oak fQ. macrocarpa) seem to be preferred to those of the black oaks*, though the bitter acorns of the black oak (Q. velu- tina), as well as of the scarlet (Q. coccina). pin iQ. palustris) and particularly the scrub, (Q. ilicijoUa) oaks are taken in increasing amounts towards the southern part of the grouse range."' "'. The red oak (Q. horealis) acorn is so large that it is seldom swallowed whole. The Investigation has two records of the meat minus the shell being eaten. Grouse apparently enjoy eating tough leaves occasionally such as those of oak for three of the birds examined had included these in their diet. One collected July 21 made almost its entire meal on the leaves of the bur oak (Q. macrocarpa). even though insects and rasp- berries were abundant. One wonders how the grouse manage to break off the stubby, tight-twigged oak buds, but they were found sparingly in the crops of eight birds from ,'\pril 3 to June 5. The Heath Family. Wherever acid soils abound within the grouse range, plants of the heath family (Ericaceae) play an important part iti tin- life of the grouse. In the Northeast, begimiiiig in July, blueberries (J'acciiiium ) and later huckleberries fCayltissacia) are much relished b\ old and young alike. Judd found .'^IM) ln-rries of / . penrisylvanicum in the crop of a September bird from Chateaugay, N. Y. Farther south they continue to be sought even in late fall as a Rlacksburg. Va.. bird killed D.-ceinber 2. had made the bulk of its last meal of 295 fruits and an additional 99 .seeds of blui'licrrirs. FrequentK large numbers of the small compact buds are taken, particular!) in u inter, for blueberries are widely distributed in clearings and as an understory in the more open woodlands. Buds, leaves or fruits of * Acrording to Wanio and Forbr*^* llic [irr urnib oak 6.S5. and while oak 2.91. of tannin in llif (ri-.li niilA from i-r.il i«|ir.'if!i arc a» follows: ri*d oak 6.01. DIET 203 these two genera were eaten by no fewer than 80 of the birds collected by the Investigation. From southern New York southward, the mountain laurel (Kalmia lati folia) supplies food, and even more importantly, shelter. The fullest crop examined, which was collected March 15, 1936, near Middletown, N. Y., was stuffed with laurel leaves, buds and twigs. Again, tough leaves seemed to be attractive. Weed and Dearborn'"' found 12 leaves of sheep laurel (Kalmia angustijoUa) in a Janu- ary-killed bird. The buds of a cousin, the azalea (Rhododendron nudijlorum) are sometimes eaten in fair quantities. Another evergreen which, because of its small size is often covered by snow, is the aro- matic wintergreen (Gaultheria procumbens), whose chewy leaves and solid-fleshed fruit are favorites with grouse. Collected near Blacksburg, Va., on ISovember 25, one bird was found by Smyth"" to have eaten enough fruits of this species to furnish 2,499 seeds. But so small were they as to represent but 10 per cent of the gizzard contents. Ofttimes associated with wintergreen is the shinleaf (Pyrola), whose flat-clustered, basal leaves are picked up possi- bly because they also are evergreen. Evergreen winter browse is furnished largely by conifers, ferns and heathers. Where avail- able, the last group is usually patronized more commonly than either of the others. The Sumach Family. — In New York, the fruits of sumachs ( Anacardiaceae) are well up in the list of favored foods. Yet difficulties experienced in cracking their hard-shelled seeds which lie just beneath the well known red coat of most of them pose a question not easy to answer. Beer and Tidyman" have suggested that such resistant seeds are taken in place of grit, a possibility discussed elsewhere in this chapter. In these days of vitamin-consciousness one wonders if some nutritive need may not be locked up in the hirsute and acid outer cov- ering of the sumach seeds. It is difficult to believe that these thin-pulped fruits would be taken merely for stuffing with so many easily obtained and nutritious substitutes usually at hand. It is a fact, however, that sumach fruits are often eaten in large quantities when an abundant supply of other foods is available. For instance, one bird collected near Ithaca, N. Y., on November 13, had swallowed 1.025 of the fuzzy fruits of staghorn sumach (Rhus typhina) along with small amounts of several other foods. In winter, one occasionally finds an astonishing amount of these sizeable seeds in a crop; one contained 1,069 seeds. Whatever the reason for their patronage, few of the sumachs are overlooked save possibly in summer. Even the poison ivy (R. toxicodendron) berry had been eaten by ten of the birds examined. Sumach buds were found but once. The Sedge Family. In the order of importance as producers of grouse food, the next most patronized group is the sedge family (Cyperaceae) . They receive greatest attention in 204 FOOD HABITS AM) REQUIREMENTS the summer, especially from the young birds. Many seeds of one genus, Carex, have been reported eaten by individual birds. Apparently tlie broods during dry seasons often fre- quent the raoister spots where the vegetation remains luxurious. Here too are found many of the sedges. As early as June 16, a chick collected in the Catskills had eaten 269 seeds representing six species. July records include those of crops from the Adirondacks filled with 1,800 seeds, from the Southern Tier with 1,515, and from the Catskills with 2,200. One adult taken in the middle of July had stripped some 4,200 seeds from Carex crinila as well as a few from three other species. The large-seeded bladder sedge fC. intumescens), eaten by 101 birds, is also worthy of mention, as is the slender-stalked sedge (C. debilis, var. rudgei), fed upon by 119 grouse. The Honeysuckle Family. Some niav be surprised that the widely-scattered viburnums and dogwoods do not have a more prominent place among grouse foods. The former plants be- long to the honeysuckle family (Capriofoliaceae) along with the little-touched snowberries ( Symphoricarpos) and the elderberries (Sambucus). Both the early fruiting, red-berried elder (5. racemosa) that loves the woods edges, and its more open-growing cousin, the com- mon elderberry (S. canadensis) however, have fruit that is well off the ground at a time when other food is abundant and for both reasons may be little attractive to grouse. Com- pared with other birds, the grouse, though favoring the red-berried elder, does not eat as much of the fruit of either species as might be expected. It is the viburnums that are the real source of grouse food in this family. Among fall fruits they rank seventh in importance, just three steps ahead of the dogwoods. The fruits of all native species are taken, though usually in small amounts. The maple-leaved vibur- num (Viburnum aceriloliuni), a woodland species, provided 74 fruits and seeds for one bird collected in Schoharie County (N.Y.) on October 5. The fruits of moisture-loving vi- burnums also are not overlooked, as is evidenced by a bird collected on December 28 from the Red River, a wilderness area jn Hamilton County (N.Y.) which had swallowed 221 fruits of the withe-rod or buckbrush ^F. cassinoides) The brilliant red fruits of the highbush cran- i)erry (I', opulus) are but slightly less attractive to grouse. Two Adirondack birds, one killed October 29, the other a month later, had swallowed 57 and 83 of these fruits, respectively. A really ambitious bird, found near Klniira (N.Y.) on November 5. had managed to collect 20 red fruits of liighbush cranberry, 42 blue fruits of arrow-wood (V. denlatuni), 25 brown beechnuts and nine other items. The Maple Family. It is surprising to find the maple family (Aceraceae) important as a source of food largely in the summer. Then two of the smaller maples, mountain (Acer spicalum) and striped (A. penns\lvanuuiii ) . frequenters of the undcrstor\ of the open woods, are sought for ilicii IdiKiiisu iuticd fruits. Those of the larger red maple (A. rubrum) are .sometimes eaten, hut more oflcn this s|)C(ics sor\cs as a source of buds, a substantial mim- ber of which are occasionallx picked up. i5uds of the well-known sugar maple (A. .<:accha- rum) rank well ahead of liiose of the oilier nienilicrs of the f;iinil\ as a source of food during the colder portions of the year. The Composite Family. The largest family of flowering plants is the aster, or composite (Compositae ) . It is entirely logical then that some members of this group should form an integral part of the grouse diet. From late fall to early spring the birds find them a con- venient source of leafy food, the most satisfactory being the hawkweeds ( Hieracium ) and the pussy's-toes ( Aniciuiaria). Occasionally a dried, long-dead leaf of goldenrod (SoUdago) or aster (Aster) may be picked iqi. The fuzz)' pappus-covered seeds characteristic of most SPRING GROUSE FOODS Trembling aspen Yellow birch Wihl black cherry Hop-hornbeam Apple Staghorn sumach Laurel Shadbush Strawberry Red oak 206 lOOl) II ABIT a AND REQUIREMENTS of the species of this faiiiih arc rarcl) taken. Tlie larger, more meaty seeds of ragweed (Ambrosia artemisiijolia), an excellent pheasant food in New York, provided an entire meal for one grouse, 1,525 heing counted. The Dogwood Family. The dogwood family iCornaceae) supplies fall foods chiefly, al- though leaves and buds from species belonging to it are taken on occasion at other times of the year. Fruits of this group furnish the tenth ranking fall food, being taken consistently but usually in small amounts. Like the viburnums, dogwoods are largely pioneer shrubs, most common in hedgerows and forest edges, although some are to be found in the more open second-growth woodlands. The fruits of one of these, the bunchberry or dwarf cornel (Cor- nus canadensis) is readily taken. Throughout the State fruits of the panicled dogwood (C. paniculata) are the next most commonly sought food in the late fall and early winter. Un- like most dogwoods which drop their fruits early, thickets of this species ofttimes yield some seeds into March. Only two records stand out with respect to quantity consumed. One is of an Adirondack bird which had picked up 182 fruits of that small conifer associate, the bunchberry; the other is of a grouse from Delaware County, (N. Y.), which, on December 20, had eaten 226 of the light blue fruits of the much larger round-leaved dogwood (C. rugosa). Though our New York records do not indicate it to be widely distributed or much taken. Weed and Dearborn''*'' believe that whenever the sour gum (Nyssa sylvatica), is common, its blue-black acid fruits are relished by grouse. The Fern Family. Fern fiddleheads are among the signs of spring. Many of the species of the fern family (Polypodiaceae), in addition, stay green all winter. The grouse, seemingly always on the lookout for leafy items, manifest a peculiar liking for the late fronds of such species as the evergreen wood ferns (Dryopteris s/iinulosa and D. marginalis). Such tough- leaved species as the Christmas fern (Polysdchum acrostichoides) and the |)olypody (Paly- podium virginianum) are also occasionally sampled. Only when matted down and covered up by snow, do the grouse cease feeding on these, to return again as the evergreen fronds are revealed by winter thaws. As spring brings out the fern fiddles they, too, receive a share of attention. In fact, the wood ferns rate thirteenth in bulk among the foods of spring, fif- teenth in fall, and thirteenth again in winter, a position of importance hitherto apparently unrealized. The Bucknheat Family. Tasty leaves are supplied by the sheep sorrel (Riimex acetosella). a member of the buckwheat family I I'olygonaceaol . Though taken throughout the year, they are of greatest importance in the spring, at which time they rank fmnteenth among the foods of that season. Unlike other members of the family, the seeds are rarely touched. The knotweeds (Polygonum), to the contrary, provide seeds only, there being no record of their leaves having been eaten. Taken usual!) in small amounts, remains of one or more seeds were found in 83 grouse. The Saxifrage Family. To the list of carl\ spring food sources should be added the members of the saxifrage family ( Saxijragaceac). The foamflower (Tiarclia cordijolia), the trim Bishop's cap (Mitella diphylla), and the saxifrages (Saxijraga) all contribute leafy bulk to the diet. Few grouse stooped to the arduous task of picking the tiny seeds. The Grape Family. In marked contrast to the taste for grapes ( Vilaceae) exhibited by grouse in some jiarts of their range, in New York they are taken in smaller quantities than one might expect. Largely because one species, the frost grape (Vitis vulpina) clings to the Raspberry Choke cherry Dogwood SUMMER GROUSE FOODS Trembling aspen Strawberry Jewel-weed Partridge-berry Sedge Mountain maple Thornapple 208 FOOD HABITS AM) REQUIREMENTS vine long after fruits of other species have vanished, the family ranks eleventh on the list of important fall food producers. Only two birds consumed large amounts. One, shot October 18, had eaten 64 fruits of the frost grape along with 58 seeds of witch-hazel (Hamainelis virginiana). Another, collected near Canandaigua in the dead of winter had a real feast. Counted in its crop and gizzard were 135 frost grapes, as well as seven other plants and four wasps. Little attention is usuallv paid to the Virginia creeper ( Parthenocissus quinquefolia) which may drop its smaller blue berries from October 'till February. The Grass Family. The seeds, and occasionally the blades, of grasses (Gramineae) are not overlooked. They supply nourishment largely during the summer months. Best liked are the manna grasses { Glycerin). Seeds of G. striata numbering 4,0.50 were found in the crop of one bird collected July 28. The reed canary grass (Phalaris arundinacea) also supplies seeds that are well received, as do the blue grasses (Poa). Blades of the latter are occasion- ally taken. In fact, grass blades furnish a small but constant source of food throughout the year. The Buttercup Family. Much earlier in the season, perhaps in late spring or early sum- mer, some of the better known spring flowers, belonging to the buttercup family (Ranun- culaceae), come in for their share of attention, especially from the chicks. Seeds of the buttercups (Ranunculus) are prime favorites, while the leaves, flowers and sometimes the seeds of the liverleaf (Hepalica) and the dainty anemones (Anemone) are taken. In the swamps, under the hemlocks and along old stumps they find the goldthread (Coptis trijolia) and sample its seeds. The Madder Family. There is one plant that has been so closely associated with the grouse as to be named after it — the partridge-berry (Mitchella repens). It belongs to the madder family (Rubiaceae), of which the only other members known to be patronized are the bed- straws (Galium). The small evergreen leaves and pert twin berries certainly must be at- tractive, for one out of every five of the "patridges" examined had been feeding on the plant named after them, most frequently on the fruits. However, only small quantities seem to be taken at a time, and they fail to rank high as a source of food at any season except sum- mer. But in the minds of most old hunters partridge-berries and partridge woods are so closely associated as to habitat that the question of how much of the plant itself is really sought out is. after all, not of major importance. The Legume Family. Members of the legume family (Leguminosae) furnish some of our best bird feed. Where abundant, as on Michigan woods roads, while clover (Trijolium repens) is relished in the fall. The birds also lake red clover (T. pralense) when it occurs in their coverts"". Best liked of all the clovers in Wisconsin is the widely naturalized alsike (T. hy- hridunij which, according to Chaddock"' makes up 38 per cent of the fall food. The grouse, however, do not care nmch for the tick trefoils ( Dcsniodium ), the fruits of which often stick in the hunter's clothing. Likewise, the hard-shelled hliuk locust ( Robinia pseudo-acacia) seeds appear to be but seldom eaten. The Jewel-Weed Family. Seeds, and rarely the leaves, of the touch-me-not (Impatiens hijlora) are the fourth most favored August chick food and rank eighth with the summer adults. They are members of the jewel-weed family ( Balsaminaceae), found commonly on central and southern New York's moist, shaly soils. These, then, are the 20 most patronized ])lant families. But there are a number of others which under the right conditions may supply food more frequently than those of some higher Tbornapple Apple Viburnum FALL GROUSE FOODS Strawberry Beech Pin cherry Staghorn sumach Yellow birch Hop-hornbeam Dogwood 210 FOOD HABITS A\ I) ItlAjVIREMENTS in average rank. For example, leaves, and to a lesser exlenl the siiiall seeds, of wood sorrel (Oxalis), are taken tliroughout the year. Particularly favored hideaways of the grouse are beneath low-sweeping evergreen branches or well up in the crowns of pines, hemlocks or spruces. Here, particularly in winter, they pick an occasional needle or even a bud or two. iNow and then they may take a cropful, as did one Greene County (IS. Y.) bird, which divided its attention on a late January day be- tween white pine (Pinus Slrobus) needles and thornapples. The needles are eaten consis- tently but in small quantities by grouse in captivity, either as a substitute for other greens or to furnish roughage. Curiously enough, pine seeds, though palatable to many birds and available in quantity every two or three years, are largely ignored. Hemlock (Tsiiga canadensis) needles likewise are eaten, though the seeds may be better liked, judging from several records. One bird taken in the western Catskills, early in March laboriously picked up 3,200 seeds and 138 needles. With little else to do from December to March, four other birds had each consumed on the day they were collected more than 1,000 seeds, the highest record being Sj.'iOO which still made up less than one-fourth of the crop contents. Included also were 244 buds of shadbush, as well as 566 buds and a large quan- tity of the leaves of the laurel. In the Northwest, the gray ruffed grouse ( B. ii. umbelloides), though occasionally making a meal of spruce buds"'", never feeds upon them so extensively as does its near relative the spruce grouse (Canachiles canadensis). Perhaps because certain members are so generally distributed throughout the grouse woods of the Northeast, the lily family (Liliaceae) is worthy of mention. Certainly the 145 adults and 16 chicks that fed on the berries of the two-leaved Solomon's seal ( Maianthemum cana- dense) found them useful. Farther south the fruits and the partly persistent leaves of green- briers (Smilax) are sometimes important fall and winter foods"'. The seed-filled capsules of violets iViolaceael are commonly taken by chicks, though they are too small to bulk large in the adult diet. The tender stalks of beechdrops (Epijagus vir- giniana) are taken by adults in late summer and in the fall. They belong to the broomrape family (Orobanchaceae). One bird started the New Year by swallowing 18 of the thick-fleshed fruits of the skunk cabbage (Syniplocarpus joclidus), an early spring favorite of grouse and pheasants alike in low swampy grounds. Allen" found mulberries (Morus) to be much relished by young grouse raised in ca|)tivity. Along with a host of other birds, wild grouse also enjoy this fruit, particularly in the South. Loomis' '. writing of South Carolina "pheasants" says, "At this season (June) the mulberry trees are in fruit, and are much resorted to. Two of these trees at the foot of Ml. Pinnacle were visited daily by pheasants during my stay." Fruits of the bittersweet nightshade ( Solaniun Ditlcaniara) liang on well into the winter in many gain<' coverts and ar<' much liked by ])heasanls. Nine nialure grouse and one chick sam- pled them. To one DcdMnlicr liird from Oswego County (N.Y.) they must have been really attractive, for it had eaten in excess of 300, as represented by more than 3.000 seeds in ils giz- zard. Another red berr\. less noticed by grouse than one might expect, is the swam])-loving winterberry (Ilex verliiillnhi) . Though it is widely (lislributcij and ils scarlet fruits persist nnlil s]iring, only eight grouse had sampled it, all in small i|niuililics. Lillle mention has been made of the various cullivated grains, for grouse survive compe- tently without liicm. In cajitivilv little (lifliculty is encountered in teaching these birds to eat cereals, including corn, wheat, oats and buckwheat. Except in cold weather when corn is a i CtAVT WINTER GROUSE FOODS Choke cherry Trembling aspen Yellnw birch Sbadbush Smooth sumach Barren strawberry Hornbeam Hop-hornbeam Apple Hazelnut 212 FOOD HABITS A\D REQUIREMEXTS favorite, buckwheat and wheat are generally best accepted, although individual grouse differ markedly in this respect. Occasionally birds in the wild may pick up one or another of the (■ulti\ated grains. One bird consumed 123 kernels of corn, though from the location it is likeh that the grain had been scattered as emergency feed for ring-necked pheasants. Eight other birds each sampled a few kernels. Grinnell'" mentions a "vivid recollection of a part- ridge which one winter could be started three times a day at the head of a small ])ond where corn had been scattered to attract black ducks'". In the Wisconsin study of fall grouse foods. Chaddock found corn, like clover, to be important. Among the items, upon which, to our surprise, none of the 1,633 grouse examined had fed. are seeds of the ashes (Fraxinus), common trees of many northeastern habitats, and buds and fruits of sassafras (Sassafras variijolium) . another locally abundant plant. The elms (Ulmus), among the producers of early flowers and fruits of spring, were generally disre- garded, though one chick did sample a seed and a single adult the swelling buds. Alders (Almis), also present in many grouse coverts, were largely ignored bv the birds. Only six grouse took small quantities of the seeds and two others ate a few buds. That the birds here draw a line is evident, but the reason therefor is not vet apparent. Animal Foods So far as bulk is concerned, animals supply a relativelv small proportion of the normal grouse diet, outranking plants only in the first few weeks of the birds" existence. Consisting largely of insects and their relatives, they represent an amazing number of forms, seemingly limited large- ly by availability. Altogether 580 different identifications of animal foods were obtained. Due to the mangled condition in which they were found in stomachs, manv could not be deter- mined as to species or even to genus. More than half again as many kinds of animals were noted as plants, but despite numerical superiority they formed only 1.1 per cent of the diet (table 25). Distributed among 165 families, they represented 11 distinct classes. Few- were of sufficient importance or taken with enough regularity, however, to warrant separate discus- sion. Grouse undoubtedly render some service to man in the destruction of insect pest«. Control of insect damage in grouse habitat is so difficult and costly that it is seldom attempted by man. Hence, the frequent appearance of harmful species in the grouse diet, even though in .*mall numbers, is worthy of mention. Such serious pests as saw-flies, weevils (such as strawberrv weevils, the black vine weevil and the popular borer), leaf beetles (including the Colorado potato beetle, the cucumber beetle and the elm leaf beetle), caterpillars (like the notorious cut-worms, maple-tree worms, canker-worms and apple-tree worms), and grasshoppers are all devoured. Plant bugs and leaf-hoppers are also taken on occasion*. Not only did the birds as a whole take a wide variety of insects, but individuals. ])articu- larly juveniles, often had picked up 20 or more different kinds in a single feeding period. One adult made a veritable biological survcv of its habitat in Tompkins Countv (N.Y.) on June 21, in consuming 27 different animals and 17 species of plants. As is true with plants, a single species of animal may provide food in different forms; the inunature as well as the adult stages of insects being taken. The egg sacs of spiders apparentlv are devoured when- ever discovered. Ants, wasps, saw-flies and their relatives ( 1I\ nu'iinptcra I are among the best sources of animal food for grouse. Altogether 84 representatives of this group were identified. * For a complete 1i«l of all inaccu ealen. lee table 173 in llic Appendix. DIET 213 many, however, but once or twice during the Investigation. Ants (Formicidael were noted more frequently than any other kind of animal food. The large black carpenter ant (Camponotus herculeanus) was eaten by 327 chicks and 168 adults, the highest incidence for any one animal. These ants were equally popular throughout the State, although the number taken was not large. A bird from Delaware County (N.Y.) ate 19 and several others selected above 15, but fewer were usually taken. The little brown wood ants of the genus Myrmica were often eaten in great numbers. A chick, on July 15, picked up 208 at a single feeding. It, together with three others, all mem- bers of the same brood, accounted for 49.3 myrmicas. Ants of the genus Formica and the little brownish cornfield ant (Lasiiis nigerj were also relished. Saw-flies (Tenthredinidae) were the second best-liked group of Hymenoplera, these wood- land pests being taken by 139 chicks and 28 mature grouse. Both adults and larvae were avidly eaten, as many as 24 of the former being found in the last meal of a young bird from Sullivan County (N.Y.). Ichneumon-flies (Ichneumonidae) also are commonly eaten. Beetles (Coleoptera) represented the most diversified food group. 228 different kinds being noted. Usually adults, but occasionally larvae, and e\en eggs, are ingested. The stomach of one bird contained 300 beetle eggs. Ground inhabiting larvae, as wire-worms (Elateridae) and May beetles fPhyllophaga) are occasionally turned up and devoured. Adult beetles are usually taken one or two at a time, but some active chicks managed to corral as many as 15 or more of the same species. One from Jefferson County (N.Y.) had eaten 18 strawberry weevils ( Brachyrhinus rugifrons) and one from Albany County (N.Y.) 15 weevils of a closely allied genus. Weevils (Curculionidae I are a usual source of food to young and old alike, those of a single genus having been taken bv 1.59 chicks. Ground beetles (Carabidae) and leaf beetles (Chrysomelidae) also were relished. Members of the genus Galenicella, American counterparts of the destructive European elm leaf beetle long estab- lished in this country, were frequently taken, as many as 14 being found in a single stomach. Moths and rarely butterflies (Lepidoptera), are eaten by the grouse during their periods of availability. Larvae are preferred, but eggs, cocoons and adults are occasionally taken. The stomach of a very young bird collected on June 10 not far from Ithaca contained 38 larvae. In a bird from Delaware County (N.Y.) 153 moth eggs were counted. Grasshoppers and crickets (Orthoptera), abundant in late summer, provide a good source of bulk food. Judd"° reported grouse from New Hampshire, in September, 1898, as feeding extensively on grasshoppers. The remains of 15 red-legged grasshoppers (Melanoplus femur- rubrum) were present in the gizzard of a bird from the Southern Tier (N.Y.). This was one of 12 chicks collected there the first three days of July which together had consumed 39 grasshoppers. As a group these 12 birds ate 243 insects of a immber of different species in addition to 15,150 seeds. If that fairly represents one meal for a brood, the total food con- sumed during the entire summer must be stupendous. Field crickets (Gryllus assimilis) also apparently are fair game for the grouse, as many as 20 being found in a single bird. Spiders and daddy long-legs (Arachnida) are eaten regularly, but rarely in quantity, evi- dently being taken in haphazard feeding rather than by persistent search. That they were found in almost two-thirds of all chick stomachs, however, testifies as to their acceptability. Stink-l>ugs of the genus fEuschistus) are more frequentlv taken than other true bugs (Hemi- |)tera), and the scale insects and plant lice (Homoptera) also received some attention. Snails are an occasional food, sometimes taken in large numbers. Almost the entire crop _>i ; FOOD HABITS AND REQUIREMENTS contents of a bird from Lewis Countv iN.Y.) was supplied by 87 pond snails I Lymnaea col- umella). Most of the records, however, are of the common land forms found under dead leaves, in rotting logs and among other debris. It is apparent from field observation that there exists at the time when most needed a wide xarietv in the seemingly unlimited supplv of insect food. They are apparently most abundant in the cover types most commonly frequented by grouse in the late spring and summer. An intensive studv of insect population by the sanijilp ])lot method in each cover type repre- sented on the Adirondack and the Connecticut Hill study areas was carried on in 1936. The investigation was repeated on the latter area the following year. In June 1936 the average number of insects available to grouse chicks was estimated at 326.()(K) per acre in the Adiron- dack study unit and 2i]0.000 on Connecticut Hill. On the latter area at least 334,000 to the acre were believed to be present the following vear. The increase was ascribed to a wetter sea- son. A more complete synopsis of the report on insect foods available to grouse chicks is given in the Appendix fp. 775 I . Seasonal Differences in Food Habits While seasonal differences in grouse food are sometimes marked, upon analysis they rarely have fundamental significance. Plant foods are dominant throughout the year I figure 161. The 100 90 70 5 :3 o > ^ 50 CD u> 30 20 10 1 1 1 • • • • • • • . • • • • • • • • • • • • • • • • • • • • •- • • • • • • • • • - • • • • ■ • • • • • • • • • • • • • • • • • « • • • • • • • • • • • • • • • • • • • « ■■ • • • • • • • • • • • • • • • • • • « '••':'•':[':■ ::■'•■'.■■ Leaves .' • ••••• • — • • • • • • ••••••• • •••••••< • •••••••• • ••••••• • ••••••• .,• • • • • • * al .",'-.'-.•. ^ • • • • • ». • • • ■ • • — • • • • • • « ■.'■■■.■•VV-VV/.v..--. .• • ^^^ . Insects 90 80 70 U) 50 J.0 30 SPRING SUMMER FALL WINTER FIGURE 16. FOOD (.KOI PS TAKEN in l.(W3 ADULT GROl SE AT VARIOUS SEASONS great bulk is produced, season by season, by a do/en or so groups, such as the .<) buds. large- toothcil rherry '>.7 buds slender-stalked fruits Birch 13.7 pin bladder yellow black black sickle Cherry 6.6 fruits. chokr bristle-stalked pale piu black buds Hop-hornbeam ... 11.2 buds. lIo|)-li(irribt>am ."..;j buds, ratkins Cherry H.2 fruits choke catkins pin llop-hornbeuni. .... 6.0 buds. ShadbushA 3,8 buds Apple :j.y calkins choke Sumach 3,6 Sumach ;i.i fruits \ iburnum 0..1 fruits staehorn stflghori) Strawberry 7.6 fruit.s highbush cranberry smooth smooth maple-leaved (Iwnrf Maple striped 3.6 seeds withe-rod .\pplc 3.5 buds Luurel :{ I leaves. mountain Strawberry .1.5 leaves mountain buds red Sumach 3.3 fruits Barren strawberry. 2.6 leaves .SlmdbusliA ... :t.o buds Dogwood bunchberry 2..> fruits staghorn smooth Hornbeam 2.3 seeds. Strawberry 2.8 leaves panicle -J Oak 2.« acorns. Ji'wel-wi-i^l '2.2 seeds buncliberry beaked red buds spotted panicleil Ainericuii black A pale ThornappleA- . , 2.2 fruits silky ml OSHT Part ridge- berry 1.4 fruits *Scientiflc names will be found in tabic 173 in the .\ppendix. aSpecicsnot identified. 216 K)()l) HABITS A\n REQl'IREME^TS SPRING FALL WINTER FIGURE 17. SEASONAL \ Mil \TI()N IN THE AMOLNT CONSL'MED OF SOME IMPORTANT FOODS OF 1.093 ADULT GROUSE IN NEW YORK >U|i|>l\. liiid' anil ciilkiii^ nl womK |)l;iiils arc still iin|)(ir laiil il<"nis of fond I hi; u re I7l. 1 he smiiiiirr diet is one nf fircat prnfiisiipii in wliirli leaves and frnils |ired(iininale. In- secl.s at this seasdii are everywhere availalde Iml tiie\ do nol iiulk lar^e e\ee|)l in the fond of the voiirifi cliieks. I Ik' fall i- ,1 lime ol Iniit liar\e.st for the firoiise with lliorna|)|)les jiiakinj; llie oulsland- lug eonlriliMlion. In anliiinn. loo. luasl is of real irn|)orlanre. e\en lhouf;li nol rejiiilarK available every year. Winter is the lime when the j;rouse turn lo luids and seeds, a diet (ontinned until early spriii-. DIET 217 Regional Variations in Foods Eaten A grouse, could it traverse all the wild lands of New York State, would find a great var- iety of habitats. The partly sandy, partly granitic Adirondack Mountain region contains coverts in many essentials representative of much ruffed grouse range in the Northeast. The high-dunied, sedimentary-rocked Catskill Mountains in the warmer southeastern part present an equally distinct set of conditions, in many respects comparable to those of por- tions of the Appalachians. The surrounding regions, together with the Allegheny Plateau, stretching westward across the southern and southwestern part of the State, represent a third type of habitat distinct from the others in important respects*. The variations (table 28) between the plant foods that the grouse take in these three regions, even at the same season of the year, are rather surprising. Differences in weather and in plant dislriluilion, of course, play their part, but it is not always easy to ascribe all the shifts in jireference to these factors alone. However, the grouse show a distinct tendency to concentrate their feeding on certain favorites at each season. These are largely the same throughout the State, except in the Catskills. This difference is puzzling, for the most-used groups, the poplars, birches, thorn- apples and hop-hornbeams, are well distributed throughout all three regions. A partial explanation for a dietarv difference in the Adirondacks may lie in the fact that deep snow covers most of the low-growing foods throughout the winter and well into the spring. The birds are accordingly forced to concentrate on the buds of a comparatively few spec- ies. The same situation holds to a lesser extent throughout the State, though this influence is offset in most regions by an earlier spring. In the Adirondacks en Sedge Mnple Cherry Cherry Cherry Sirawberry Summer Do p wood Thornapple Maple Jewel-weed Sedge Thornapplc Dogwood Jew el -weed Strawberry Violet Partridge-berry Shadbush Beechdrops Viburimm Buttercup Buttercup Wintergreon Birct. Apple Beech ThoriiappN' Hop-hornbeam Thornapple Cherry Thornapple \'iburnuni Viburnum Strawberry Apple Fall Hawkweed Birch H(i|i-hornlMMitn Wood sorrel Cherry Cherry Hop-honibiMiM Sumach Sumach Laurel GraiK- Wood fern Clover Dogwood Oak niij'W(K)d Bee<:li Birch Cherry Hop-hornbeam Aspen Birch Birch Cherry Hop-hornbeiirn Barren striivvlM-rrv Birch As[K!n Shadbush Sumach WiiiUT Hazelnut Cherry liop-hnrnlH-iim Shadbush Laun-I American hornbeam Apple Strawberry Apple Wood fern Aspen Barrc!! slniwUTry Viburnum Sumach 'rh(>rna|»|»Ie Maplo W illow Yearly \'akiations in Consumption di- Foon Items A conipai Imiii oI ihc fnods taken bv grouse over an 11-vear period brings to light some interesting inconsistencies (table 29). In glancing over this table, one might conclude that the birds exhibit little conlinuitv of taste year bv \car. This is especially pu/.zling wlicn the parts eaten arc largely buds or calkins, as in the asi)cns and the hop-hornbeam, the quan- tities of which, available vcar bv vear, apparcnih arc iniicb llic same. Another grou[) in DIET 219 which buds and calkins are also the chief parts eaten is utilized fairly consistently. In this are found the birches and maples. Then there is a third group, largely fruit-producers, such as cherries, sumachs, strawberries, sedges and blackberries, which exhibit inconsistent and substantial fluctuations in use by grouse over the 11 years. TABLE 29. YEARLY VARIATIONS IN PERCENTAGE BULK OF IMPORTANT FOOD PLANTS OF ADULT GROUSE IN NEW YORK IN COMPARISON WITH THE 11 YEAR AVERAGE Yeats Average 1931-1941 1931-32* 1933 1934 1935 1936 1937-38* 1939 1940 1941 12.4 10.6 9.2 8.8 5.7 4.5 3.8 3.8 3.3 2.7 2.5 2.3 2.0 2.0 1.6 21.8 1,093 1.6 7.4 11.9 1.5 12.7 9.9 3.7 4.2 0.8 2.4 A 0.2 0.9 1.4 2.8 38.6 53 15.3 10.0 11.6 3.9 7.6 3.1 2.9 2.9 0.4 3.3 ... .A 0.7 0.2 1.4 2.2 34.5 94 13.0 6.6 10.8 2.8 9.2 2.8 1.0 4.3 8.3 0.7 2.0 0.2 2.0 1.1 1.0 3t.2 164 17.3 17.7 10.5 1.6 1.7 4.1 3.3 6.5 1.9 4.3 4.0 0.4 2.4 0.7 1.2 22.4 302 8.5 12.1 13.6 3.5 10.4 3.0 2.5 3.1 0.2 4.0 4.1 0.8 0.1 0.7 0.4 33.0 111 6.7 6.8 17.8 1.1 10.0 5.2 2.8 0.6 8.5 2.6 1.3 0.8 5.2 0.5 1.6 28.5 97 5.5 5.4 9.7 13.4 5.3 r* 2.3 3.2 3.8 4.0 4.2 0.2 0.7 0.8 22.2 88 22 9 2()!l 9.4 4.9 10.7 4.2 2.4 0.9 2.5 2.1 2.4 3.1 2.3 3.6 0.6 7.9 97 25.9 Cherry 8.7 Birch 10.3 Raspberry, etc I lop-hornbeam .... Thornapple Strawberry 8.8 7.0 1.2 0.7 0.2 Beech 1.0 2.2 Shadbush Sedge 3.1 1.9 O.S Maple 3.6 Dogwood AU other foods No. birds examineii. 3.8 21.1 57 *Coaibiaed, since the number present in a single year was too fewatu tabulate satisfactorily. A Not recorded. Reasons for tliese variations are interesting but certainly still conjectural. Precipitation and temperature conditions are probably basically responsible for fluctuations in the abun- dance of many fruits. A shift in the carbon-nitrogen relationship is said to control a cycle of approximately three years in the occurrence of beech mast. In years of plenty, the nuts are eaten in large quantities, their very size being conducive to their bulking large in the birds' diet. In non-beechnut years, however, the grouse fall back on other sources of food, notably sumach, thornapple, cherry and aspen. In each of the years studied, the bulk supplied by all four of these groups together was very similar. In 1934 the consumption of beechnuts was about three times that of the subsequent two years, and products of the other three species were eaten correspondingly less that year. As yet there is no satisfactory explanation for the yearly changes in consumption of the always available buds. Though the study should be continued over a much longer period before all conclusions can be stated with certainty, it seems clear that changes in utilization are many times dictated not by availability, but rather by some other factor not yet apparent. One has but to compare the 11 -year average with that for any one of the individual years to raise the question as to the degree of importance to be assigned to studies covering short periods. The bulk of many of the foods taken in any of the 11 years here compared shows wide variations from the 11-year average. Deviations too large to be ignored are likely to be obtained by drawing conclusions on the volumes taken over short periods, unless one is in- terested primarily in obtaining a mere list of foods and a rough indication of the extent to which they are utilized. The degree of reliance that one may place on such generalities is indi- cated by the fact that, of the five genera taken in greatest volume over an 11-year period, in only four years were the same five at the head of the list, and even then the order was shifted in each case. In 1934, 1935 and again in 1939 but three of the group were among the first five. 220 FOOD HABITS AM) RE(^LIREME\TS The variety of choice also differs markedly. On an average, the 15 most used foods supplied a little over three-fourths of the volume of all food eaten. From 1931 through 1934 they dropped to less than two-thirds, while in 1940 they provided the astonishing amount of 92 per cent of the bulk for that year. Food from the birches was found most consistently, while the volume of aspen taken varied all the way from 1.6 to 25.9 per cent. Variations in Diet Throughout the Range No matter how carefully food habits research may be carried on in one region, much can be learned by comparing the results with those obtained elsewhere. This is not always possible, however, for records are occasionally gathered without reference to age of the birds or sea- TABLE 30. A COMPARISON OF THE VOLU.METllIC PERCENTAGES OK FALL FOODS CHOSEN BY ADULT GROUSE IN NEW YORK WITH THAT REPORTED FROM OTHER PARTS OF ITS HWGE* Region Foods New YorkA (Bump & Jones) New England (Gross) Pennsylvania (Kuhn) Virginia (Nelson et al) Wisconsin (Chaddock) Important in New York State: 15.9 10.3 6.4 4.6 4.5 4.2 3.9 3.8 3.3 3.1 3.1 2.4 2.3 1.9 1.8 1.7 1.2 0.5 0.5 0.5 0.4 0.3 0.3 0.3 0.3 0.2 0.2 trace trace trace 7.3 0.9 3.7 5.3 4.7 5.7 0.6 2.5 nl i'.ii 2.9 3.0 O.H 0.6 2..! ' 3.8 3.3 1.2 trace t 11.6* 4.8 2.0 4.1 3.2 0.5 3.4 9.0 1.7 4.0 2.3 0.9 6.1 1.1 11.5 ■4.7 1.8 3.8 '4.8 2.4 1.0 trace trace 0.4 0.3 9.0 0.9 2.3 0.9 10.6 1.4 1.1 0.4 O.S trace 4.9 4.2 3.9 "bil 16.3 7.9 ' 3.3 ■M Heerh 0.3 0.5 Hirch 9.5 Oak 0.6 Shudbtish 38.4 1 2 Wood forn tni|Hjrtunt elsowherc: Hoan Wiiitor^Tci'ii 4.9 Ast^ liX) JUNE JULY AUGUST FIGURE 18. AMOUNT AND KIND OF FOOD EATEN BY 540 GROUSE CHICKS DURING JUNE, JULY AND AUGUST 224 FUOU HABITS AND REQUIREMENTS Though in captivity the chicks feed freely on ap])le leaves, in the wild only a single individ- ual of all those examined had sampled them. Animals bulk largest in the June foods of chicks, constituting 56.6 per cent: drop sharply in July to 13.0 per cent and in .August to 5.0 per cent. At this time the diet approximates that of the adults. It is doubtful if chicks search niuih for insects after they are several weeks old. for they seem to concentrate on finding suitable })lants. picking up animals only inciden- tally. Running back and forth in a most haphazard manner, pausing now and then to probe beneath a leaf, it is no wonder that the chicks uncover such a variety of insect life. No bug seems too bitter nor caterpillar too ugly to be tried at least once. In the 540 chicks examined, 475 different animal foods were identified. Many ate 20 or more kinds and several upwards of 30. Protective devices are no certain deterrent, for the formidable lo moth larva, one of the fiercest-looking of all caterpillars and armed with poisonous spines, fell prey to one chick and stink-buss seem to be taken with relish. TABLE 31. MONTHLY VARIATIONS IN THE IMPORTANT PLANT AND ANIM.\L FOODS OF 540 NEW YORK GROUSE CHICKS Months K(km)s June July August Volumetric percentage Number of birds in which found* Volumetric Iiercenlape Number of birds in which found* Volumetric percentage Number of birds in which found* PlanU 6.1 1S.7 10.8 0.2 3.0 1.6 0.2 0.3 ....t traceA traceA ....t 0.8 5.0 7.3 3.1 1.3 6.2 4.0 l.S 0.4 3.7 7.6 2.0 2.6 6.5 4.8 0.6 71 142 124 9 62 30 2 2 ...t 4 1 t 26 178 88 78 44 147 129 55 41 89 108 30 95 71 105 24 42.1 16.4 6.6 4.4 3.7 3.S 3.3 1.3 0.9 0.8 0.5 traceA 0.4 4.9 0.2 0.5 0.4 a.o traceA 0.2 1 rare A o.:i 0.7 2.3 0.2 0.7 1.5 0.8 178 176 94 69 110 100 31 17 9 40 17 43 174 49 51 14 88 6.1 15 34 64 55 43 72 36 93 32 5B.4 6.6 traceA 10.6 2.2 1.1 ....t 0.7 2.8 1.4 1.6 1.8 0.1 1.9 0.1 traceA traccA traceA 0.1 0.1 traceA 0.1 0.1 1.2 traceA 0.1 0.8 0.5 IIT 128 6 8") 42 40 ...t Mnpl,. 11 28 25 22 Yew 10 15 Animnls liitiecUi: Hymt'iiopt^'iu: AiitA 136 2 16 5 Hcctira (r.oliHjpU'ra): \Ve*'vil8 Lpfif bcetU^s 37 23 6 17 Other brrili's Moths ( I^'|iier of occurreniTeH for each (trnup: many birds ale wveral repreaeiilatives of a Kroup. ATrace. or iosa than (t.l pT cent. tNot taken. tlncludes (Vnrrinitini) and (Cityltltanrint. Ants, often considered as "protected" insects, are the most poi)ular fo the SIO tabulated containing one or more of them. Larvae and adults adults of ichneumon-dies are also well liked, 139 young birds eating the latter. Beetles, largely weevils, leaf beetles and ground beetles, along w ders and grasshoppers, supplied the bulk of the remaining animal food. od. 488 stomachs of of saw-flies and the former and 1 45 the ith cateriiillars. spi- DISTRIBUTION AND HABITAT REQUIREMENTS OF GROUSE FOOD PLANTS 225 Monthly Variations in Summer Chick Foods From June through August there is a progressive change from an animal to a vegetable diet. At first the ])lant foods are few in number and consist for the most part of the smaller seeds and bits of easily reached plants. Feeding activity during the first ten days after hatching consists largely of catching ants and beetles and occasionally capturing a juicy caterpillar, interspersed with the diligent striijping of sedge seeds and sampling of strawberries. By July animals make up only a tenth of the food and the birds are patronizing a wide variety of plants, even including some fruit-bsaring trees. The consumption of blackberries, the bushes of which provide both food and shelter, rose rapidly and amounted to 42.1 per cent of the total for the month. This gain was largely at the expense of animal foods, which dropped from 56.6 per cent in June to 13.0 per cent in July. The foraging becomes still farther diversified in August and in later summer the young tend progressively to take about the same foods as do the adults. The monthly variations of the most ini|)r)rtanl |)lant and animal foods of chicks are indi- cated in table 31. DISTRIBUTION AND HABITAT UFQUIREMFNTS OF GROUSE FOOD PLANTS Plants furnishing food for grouse at one or more seasons of the year are to be found, often in abundance and \arietv in incisl overgrDwn fields and woodlands throughout the Northeast. Some, such as \aric)us species of wild cherries. as|)ens. liiaikberrie.s. raspberries, birches, sumachs and thornapples, which together supply It! per cent of all grouse food, occur in every region of the State. They thrive on almost all types of soil, from sand to clay, from rich to poor, from wet to dry. For most of them, the one common requirement ap- pears to be an absence of strong competition for growing space, for many characteristically occur in the earlier stages of plant succession, as on cutover or overgrown lands. Pin cherries, aspens, gray and black birches and sumachs are short-lived plants intolerant to shade. They build up a ground cover and prej)are the way for the more shade tolerant and longer-lived species such as black cherry, ash. yellow birch. l)eech and maple, which follow them and ultimately shade them out. Other grouse food plants of overgrown lands as apple, thornapple, shadbush and hop-hoinbeam are perhaps more resistant to shading and therefore may persist for a longer lime under the closing forest canopy. Only a few such as the mountain and the striped nia|)Ies. are t\|)i(allv understorv shrul)s which thrive in relatively dense shade. Of the many plants eaten in large (]uantities by grouse, but few are limited as to dis- tribution in New York by soil or climatic factors. Of the birches, neither black nor gray is found widely in the Adirondacks, nor is the latter conspicuous far west of the Hudson Valley. Hop-hornbeam is most abundant in central New York; less so in the mountains and on Long Island. Huckleberry, blueberry, bunchberry and wintergreen thrive on mod- erately to strongly acid soils. Partridge-berry and evergreen wood ferns reach their best development in the duff of the forest floor. In table 32 are recorded some of the habitat preferences and other characteristics of ruffed grouse food plants. Though many of the species may be widelv distributed, most are restricted more or less to certain stages in plant succession. Grouse habitats are made up of plant associations. For convenience in identification we 226 FOOD HABITS AND REQUIREMENTS divide these into cover types. These may be classified as open land: overgrown land: hard- wood, softwood and mixed forest; slashings; and hums. In New York, by far the largest part of the open land rover type, in the territory where grouse are common, is composed of poor hill pasture, together with a little meadow and crop- land. Farming here is a precarious business at best and abandoned farms are becoming in- creasingly common. When this occurs open areas revert to "poverty grass," weeds and moss. I'ASTIKI..^ AM) 111.1.1).^. 1.1.11 lu THEMSELVES, SOON REVEKT TO OVERGROWN L.\NU^. lll.UL M'H- CIES PROVIDING EXCELLENT GROUSE FOOD AND SOME SHELTER ARE APT TO BE PLENTIFUL The character and composition varies with the last farm use and the degree of soil depletion. Especially on old meadows and in better pastures, an abundance of strawberries and of sedges, both seasonallv important to grouse, may be produced. Areas that have been denuded by lire also may grow sedges, but seldom produce an abundance of strawberries. Even before abandonment, ill-kept pastures and fields may be invaded by brush species such as sumachs, briers, scrub apples and thornapples. Because of unpalalability, thorns or sheer toughness, these plants are able to endure moderate browsing by stock. With this pressure removed there develops a typical overgrown land association of the above mentioned species together with pin and choke cherries, aspens, birches and other trees. These establish themselves first in the protection of stone piles, stumps and old walls. From here they spread in ever-increasing numbers. As the density of the stand and the consequent competition for growing space increase, the tree species arc able t<■>[■» ■»■«■>[ ■¥■¥■¥* ** * **♦ M * ** **♦* ** *** *-♦(♦♦* ♦** * * M*************** ** ■K-n-tm-n-x ♦■»♦♦ ♦♦♦♦ -fc -x-tr-tc-ti-tm-n-n-tum *«** *♦ * ♦ paE[Bi aooq o-t;olai c il is an i-\ccc(lingl\ FOOD AVAILABILITY AND SHORTAGES 229 dangerous one and should be used with caution and then only by experienced operators. This item is considered in detail in Chapter XVI. FOOD AVAILABILITY AND SHORTAGES As has been noted, the list of plants patronized by grouse in New York alone includes 994 items. So widely distributed and commonly found over much of the State are many of these that there are few grouse habitats without a comparatively large native supply of food- producing plants. Because a plant is present, it does not always mean it is producing grouse food, for some do not fruit or seed except under favorable conditions. Most of the food plants are primarily pioneer or transition species which do best in the sunlight along woods edges, in overgrown lands, in cut-over areas or in blow-downs. Insects also are most abundant in well-lighted places where the vegetation is varied and lush. Grouse might not starve if they were confined exclusively to heavy woodlands, but they would not find there a majority of the foods they like best. So long as the territory of an individual grouse includes openings, whether made by man or Nature, that foster the pioneer and transitional food species, these may usually be depended upon to produce a sequence of foods which me?t the birds' needs throughout the year unless grazing or cultivation intervenes. Of course la'k of handy and suflicient shelter may make the food largely unavailable, for both cover and food must be present and adequately inter- spersed to make a grouse habitat productive. There is no evidence that grouse have ever starved to death in the Northeast. The weights of adults collected in numbers throughout every month of the year show relatively slight variations. These can be explained in part on the basis of seasonal activity, in part by the less adequate nutritional character of their winter and early spring diet. Even the heaviest sleet storm does not armor buds and twigs long enough to produce a serious food shortage. Grouse held without food fail much more rapidly than do pheasants or quail. A period of from three to four foodless davs under winter conditions produced an average loss of 22 per cent in the weight of birds kept in captivity. They had eaten only natural foods during the preceding weeks. Experiments* indicate that grouse have very little metabolic or chemical reserves fri>ni which to endure much longer periods without food. Whether they suffer from shortages of certain nutrients one cannot sa>. Helati\eU little is known as to the chemical composition of grouse diet and nothing as to the degree of physi- ological availability of the nutrients in their foods. Scientific studies of game bird nutrition, as yet in their infancy, must await the production of a large number of grouse in captivity, as well as improvement in techniques for handling them. In the meantime, in summary food items known to be taken by grouse in large quantities are available in large amounts throughout the year in every inhabited grouse cover known to the Investigation. No starved birds have ever been found. No sharp weight losses have been recorded even in adverse weather. Food shortages in the generally accepted sense are believed, therefore, not to occur. A few of the more abstract relationships of food to health are pointed out later in the chapter. EFFECT OF FOOD ON GROUSE DISTRIBUTION At some seasons grouse exhibit a marked preference for coverts where certain foods are easily obtained. Curiously enough this is most evident in the summer and fall when food is * Set- Appoiicliv, p. 7(il. 230 FOOD HABITS AND REQUIREMENTS most abundant. The drawing power of a beech ridge in a good nut year is well known to many grouse hunters. The Investigation has records of grouse being flushed from thornapple clumps well over half a mile from their usual coverts. Hedgerows with choke cherries and frost grapes ofttimes furnish the incentive for a bird or a brood to feed at a considerable dis- tance from its normal haunts. Sj)iller''" records concentrations of birds in fall feeding areas in groups too large to have been drawn from the immediate locality. So far as one can judge, these excursions may represent a temporary letting down of territorial bars for the old birds; birds of the year of course have not as yet established territories. From the first of November, or earlier, especially if there is a blanket of snow, the birds lyH- ally forsake such exposed feeding places in favor of those closer to adequate shelter. Not as pronounced, but still definite, is the tendency to frequent cut-over areas, woods edges or overgrown pastures where the vegetation is varied and luxuriant. Here are found ample amounts of easily obtainable food combined with adequate shelter. The Investigation found these areas especially attractive in summer, when they were utilized by 26 per cent of the birds in contrast with 13.9. 8..5 and 13.6 per cent for fall, winter and spring respectively (table 152, p. 818). Broods in particular are to be found in the slashings during the warmer months (table 135, p. 801). Though these may occasionally form a dense tangle, the spots most frequented may be rather open with food and cover interspersed. In fact there is a tendency to shun dense cover when not associated with a variety of other vegetation, especially in the summer and fall. This applies also to evergreen plantations. Observations do not indicate that adults need to travel far beyond the boundary of their established territories for food at any season of the year. The usual territory of a grouse in- cludes winter shelter, spring breeding grounds and summer and fall feeding coverts. If all of these are not present within easy flying radius, say one-half mile, it is not likely many grouse will set up housekeeping there until Nature or man provides the missing requirements. COMPETITION WITH OTHER SPECIES FOR FOOD Few species are less affected by competition for food, especially in winter than is the ruffed grouse. Adequate quantities of buds are always available. It is therefore no hard- ship to share the various berries, fruits and seeds, even in hard winters, with other birds and mammals. Grouse are quite independent of grains such as buckwheat and corn. A possible relationship between adverse growing seasons, resulting in a lowering of the nutritive value of grouse foods, and the health of the birds the following spring has al- ready been mentioned. If this hypothesis is ever verified, one might wish that the grouse were more keenly interested in the more nutritious cereal.* and berries in addition to buds. However, fundamental grouse feeding habits are not easily changed as was demonstrated on the Connecticut Hill area in the fall of 1932, when 240 shocks of buckwheat were placed in well occupied cover. Only one bird was found to have utilized the grain for food. Similar results have been noted in attempts to entice grouse to return to such favored fall foods as beechnuts and tliornapples made available at feeding stations after winter snows had forced them to adopt a diet consisting mainly of buds. -Ml this tends to indicate that grouse are not hard-puslioil for food even in winter and that other species of wildlife do not compete seriou.-Iv with it in this respect even at that season. THE EFFECT OF OPENING UP THE WOODLAND ON FOOD SUPPLY 231 THE EFFECT OF OPENING UP THE WOODLAND ON FOOD SUPPLY Left to her own devices. Nature eventually produces a type of vegetation termed the cli- max, which, generation after generation, succeeds itself. In New York, physical condi- tions run the gamut from seashore sands through cover of every age and density to Mt. Marcy's treeless lop. Climax conditions show a correspondingly wide variation. In woodland habitats such as are occupied by grouse, the forest climax is most often represented by a mixture of large conifers and hardwoods, with interlocking crowns shad- ing out all but a sparse vegetation beneath. In such situations food and shelter for wildlife living above ground but largely beneath the crown canopy are often scarce indeed. However, natural forces tend constantly to break up the continuity of this forest cover. Diseases such as the chestnut lilighl: insects such as bark beetles and the defoliating cater- pillars; weather, particularly late frosts, drought and wind; fire and man's axe often cause the forest to revert to earlier stages of plant succession. Carried far enough, brush takes the place of woodland, grass takes the place of brush and the plow furrows under the grass. Left alone. Nature then begins a slow process of environmental rehabilitation which, if not disturbed, ultimately will result in the reestablishment of the climax type. The ruffed grouse finds its food and shelter needs best combined in cprtnin of the in- termediate types. Few individuals are found in cultivaled areas because of ihc paucity of acceptable foods and shelter. Meadows arc likewise avoided except where they adjoin woodlands or brush areas, even thniijjh tlii-\ siip|)orl suih choice food plants as sedges, butlercups. (lovers and strawlierries, and a host of acceptable insects. \^Tiere the shrubby vanguards of the woods arc invading grasslaiuls. however, greater concenlralions of grou.se foods usually occur. It is in this zoned intermingling "f field and forest liiat wild apples and thornapplcs (ind ihoniscKcs most at home. .Stout-stemmed blackberries form prickly tan- gles and there are clumps of dogwood and viburnum, largely bird-planted, along the field bor- ders. Though blueberries and the larger-fruited huckleberries may be spreading, if the soil is acid, they have not as yet crowded out the strawberries, grasses and sedges which grouse patronize. Pioneering trees, such as aspen, cherry, and birch, whose buds or fruit the grouse relish, may also be seeding in as a prelude of forests to come. Furthermore, the number and variety of insects in this type of cover is enormous. In time, of <-oursc. the more permanent tree species take over, closing in the crown cover and eliminating the light-demanding .shrubs, aspens and cherries. The forest has reestab- lished itself. Habitat, dissimilar in appearance, but rich in grouse foods, may be created by heavy forest cutting. Here, a great variety of seeds has lain dormant in the duff ready to spring to life at the coming of light in all the verdant confusion characteristic of a cut-over area. Many brusliland species, as well as a host of herbs, shrubs and trees will thrive in such slashings. Judging from the number of grouse, and especially those with broods, which frequent such spots from June to November, food and shelter conditions here must be nearly at their best. But again as cut-over areas grow up into woods, it is inevitable that many of the plants furnishing food for grouse give way to the climax species. Under them such shade-toler- ant plants as the evergreen wood fern, partridge-berry dwarf cornel and witch hobble, all good grouse foods, gradually fill in a small part of the gap left by the shading out of the luxuriant, light-loving vegetation of the slashing. 232 FOOD HABITS AND REQUIREMENTS Such is the story in New Yoriv and throughout many parts of the Northeast. Eke- where within the range of the grouse, the plant speries characteristic of each stage in the succession may differ from those here named, hut the resuh.* in terms of productive grouse habitat seem to i)e much the same. It is well recognized that good grouse coverts must include some of these openings which seasonally produce an abundance of food and shelter. A brief consideration of how these naturally occur, or may be made by man is, therefore, not amiss. Natural Openings These are largely the result of fire, weather, insects or diseases. Except in the case of severe fires the succession is seldom set back to bare ground. When the burn is deep or repeated, however, the natural |)roductiyity of the site is usually so impaired as to make the re-establishment of desirable food and cover a matter of many years, although such species as the pin cherry, aspens and blueberries may soon seed in. Ruffed grouse, unlike the sharp-tailed grouse which habitually frequent burns, are seldom found here except along the edges, near shelter. Light burns, if continued, may have the effect of decreas- ing grouse food supply through encouraging the spread of grasses and weedy ground cover. Reduction of the more desirable food plants, together with the gradual elimination of small shelter-producing shrubs and conifers by fires, is likely so to change the habitat as to lower materially or wipe out its carrying capacity for grouse. This is |)articularly true within the woodland itself. But fire is not always detrimental. Occeisionally in a grown-up slashing a light burn may set the succession back perhaps ten to fifteen years, thus allowing many desirable species which otherwise would be .shaded out, to re-establish themselves. In fact, even a severe burn, could it be limited to a few acres, might be useful in the same way. The utilization of fire as a tool in wildlife management is discu.ssed in somewhat greater detail in Chapter XVI. However, it is at best a dangerous servant. The effect of weather in creating or maintaining openings is more wide-spread, though probably less well defined than is that of fire. Many an abandoned field in New York does not grow u|) rapidh to shrub or tree species, primarily because the alternate freezing and thawing to which it is subjected causes heaving that destroys seedlings. In wood- lands the effect of unseasonable frosts is ofttimes strikingly evident. In 1936, in the Adi- rondacks, late frosts pruned the early spring growtli (if bccrb. thus temporarily letting in more sunlight to the forest floor. Occasionally, also, extremely severe winter weather may kill le.ss resistant species, thus creating small openings which increase feeding opjiortuni- ties for game. Much more violent are the effects of wind. yar\ing all the way from breaking off or up- rooting individual trees to opening up thousands of acres of closed canopy woodlands. The destruiti\c hurricane of 1938 that swept through Long Island and \cw England is an example. Less clearly recognized is the effect of sun and air movement so drying out the soil in a fresh-made slashing as to eliminate many of the le,*s adajitable trees and shrubs left after lumbering. The result is to create or maintain small patches of light-loving |)lanls wbicli arc fiuorcil ii\. nr can adjust themselves to the new conditions. An ciTeet in nian\ wa\s -iimilar is sometimes brought about by large-scale attacks of in- sects such as the larch sawfU or of tree diseases, as (he chestnut i)liglit. Tliroughoul the (]ats- kills in llie l.ile |9.'i()"s ibe forest lent caterpillar defoliated sugar ma|)Ie> fur several years so THE EFFECT OF OPENING UP THE WOODLAND ON FOOD SUPPLY 233 completely as to cause the death of thousands of trees. Where maple was the dominant spec- ies and the undergrowth was not too dense, this resulted in many small openings, which were quickly occupied by a host of herbs, briers, sprouts and insects. The result was a rapid increase in grouse food. Thus natural forces may create many breaks in forest cover. Man-Made Openings Spurred on by economic necessity, man ma\ cau.se openings in woodlands of a yariety of forms. Each of these may differently affect the grouse food supply. The most seyere of these is of course, wholesale clearing and subsequent culti\ation. Crops such as buckwheat, corn, clover, together with accompaning weeds and insects, unless located immediately adjacent to adequate shelter are seldom of i7iuch value to grouse, as these birds rarely venture far into the open for their food. Allowed to revert, however, as is occurring in many parts of the Mortheast. cultivated fields ma\ be taken over by a |)roduclive combin- ation of trees and shrubs, both evergreen and hardwood, intergrown with gra.sses. straw- berries, clovers and other much sou":hl after herbs. FROM JUNE THROUGH SEPTEMBER NO COVER TYPE IS SO RICH IN GROUSE FOOD PLANTS AS ARE CUT-OVER LANDS (Grazing, particularly if heavv. lends to eliminate woodland undergrowth and to encour- age replacement of the normal flora of the forest floor with grass. Light grazing may be beneficial, however, where the undergrowth is dense and in need of being opened up. Many 234 FOOD HABITS AND REQUIREMENTS lightly pastured woodlands in New York produre good crops of grouse. Cattle help also to maintain edges and to keep the woodland from encroaching too rapidly on the open ground. Here also occasional catlle-sown wild apples spring up. Thornapples and certain of the dogwoods and vihurnunis, together with a sprinkling of conifers which the cattle do not like, furnish much-frequented feeding spots along the line where pasture and forest meet. Many of our finest summer and fall grouse feeding grounds in New York have been es- tablished in this way. The compacted earth of the cow paths is slow to change, but the brushy areas, with their interspersion of conifers, and open spots may provide almost per- fect feeding grounds for broods and adults from June to early November. Occasionally the grouse also choose such locations for their nests. But of all man's tools, no single one has wrought such far-reaching changes in grouse habitat as has the axe. By breaking up the forest canopy it has altered the environment so as to release light-loving trees, shrubs and herbs. In the openings thus created are to be found the greatest variety of grouse foods, both plant and animal, for lumbering not only lets in the sunlight but usually disturbs the soil as well. Thus is produced, often in the space of a few acres, the whole series of environmental conditions from bare ground to woodland. The effect is dependent in large measure upon the site, the age and compositinn of the forest before cutting, the severity of the cut and the length of time since it was done. The better the site, the more varied the forest make-up, and the more severe the cut, the greater is the chance that the cutting operation will produce a profusion of grouse foods. To understand the effects of patches of cut -over woodlands upon grouse production knowledge is required of conditions preceding as well as following the catling operations. It was, therefore, necessary to make a study of slashings under controlled conditions. This began in 1932. when a series of openings varying in character and in location were es- tablished. Subsequently 341 clear-cuttings were completed as test plots in 8 of the State's widely scattered wildlife management areas. Conditions preceding cutting and at regular intervals subsequently have been noted. In most plots detailed measurement of site changes have been carried out as discussed in the preceding chapter on Shelter. Analysis of results reveals that diversification of the vegetation may start within a nuintli after cutting. In terms of grouse food production this tendency reaches its height in from three to five years. As the cut-o\er area continues to grow up. many liglit-loving grastures or slashings, seldom contain adequate amounts of s])ecies fumisliiiig winter shelter fr)r grouse. There- FOOD AND FLAVOR 235 fore, when too extensive, they are not usually coiiduiive to maintaining large grouse populations. Many small slashings widely scattered and of different age classes are to be preferred to a few areas occupying many acres. FOOD AND FLAVOR One of the few points of general agreement among those who hunt the grouse is that its flesh is exceedingly palatable when properly cooked. Many an early author has ascribed this to a particular liking which the bird is supposed to exhibit for leaves and twigs of a pungent flavor. Eaton'" lists "wintergreen, mint, sorrel, birch and various kinds of ber- ries which impart a peculiar gamey flavor to its flesh". Maine birds, according to Mathews'^', "are particularly fond of the buds of the black birch which give their flesh a peculiar and very agreeable flavor". That this is not a new thought is proved by Alexander Wilson's'"", statement in 1812 that, "The 'pheasant" is in best order for the table in Sep- tember and October. In this season they feed chiefly on whortleberries and the little red aromatic partridge-berries, the last of which gives their flesh a peculiar delicate flavour". Additional backing for the food flavoring flesh idea, though with a diflcrcnt meaning, comes from Elliott"', who indicated that "its flesh, as is well known, is light and tender, but in late fall and winter becomes bitter because of the bird having fed on the leaves of the alder and to many persons is then (]uite poisonous". In sjieaking of Sabine's rufled grouse [li. u. sahiiii). he continues. "The flesh is white and palatable save in winter, when it is often bitter, oica.*ionally fla\ored with turpentine from eating the buds of the fir trees". In speaking of the gray rufled grouse in the Yukon Terri- tory, he adds that it subsists on spruce buds which gi\e a disagreeable flavor lo the flesh. In this he is eloquentl) borne out by Sandys'"', who writes of the grouse picturesquely, "Its favorite food is the buds of the spruce which impart to the flesh a flavor which might ap- peal to the ])alate of an eastern spruce gum-chewer, but which signally fails to hold the appreciative attention of an epicure unless he also happens to be a lost prospector keen for a 'grubstake' ". The presumably poisonous properties imparted lo tlie flesh of birds that fceil on laurel has already been discussed. The present authors are duly appreciative of this background of testimony, but being neither spruce guni-chewers nor epicures tliey ha\e failed to note ])ersonally llie peculiar flavors described by these observers. The grouse, shot in season and properly prepared, is in New York invarial)ly a delicacy of the first order. FOOD AND ITS RELATION TO HEALTH The ability of the grouse to survive the rigors of a biting winter on little more substantial foods than buds is but one of the interesting situations confronting those who would study its food habits. Pheasants and quail would starve miserably on a diet of cherry, asjjen and birch ^i^. ^'^^■x ^'i'^ "l^lS^Mn'^-i/fi: ?ifi?-^ 236 FOOD HABITS AND REQUIREMENTS buds, all of which are staples for the rufled grouse in New York from November until April. True, they eat much more in proportion to their size, for the normal contents of a grouse crop in winter is almost as large in volume as is that of a pheasant, though the latter is usually twice as heav\ . The abilitx to utilize buds for food is one reason uh\ the grouse ia:i sur\ive from Alabama to Alaska. There are other ways in which food affects the existence of the species which arc much less easy to identify, for there is much more to the study of food habits than the mere recognition of the amount of each kind which is eaten. True, in analyzing the relation of food to health, this is the first step. This done, the question naturalh arises as to the nutritive value of each food to the bird. One can not logically assume that ati item commonlv eaten, bv that token, must provide substantial nourishment. The next step, therefore, is to determine the amount of each nutritive constituent — proteins, fats, carboh\drates and minerals — that are present in the foods that constitute the normal diet. This can i)e accomplished by rather involved chem- ical analyses. But all birds are not alike in their abilitv to break d\ othc-r less accurate methods. Variations in ihc- weight of grouse in relation to the amount consumed of a single food item or of a combiiiatioTi of several kinds have been cldcriiiined experimentally with hand-raised birds at the Research Center. A start has also been made at testing the results of feeding grouse o\cr a considerable period of time on one food by checking progressive changes in their ph\ siological reactions. The first of these methods yields results which arc at best onl\ indicative, while tlic second invciKc's such liighU specialized ccpiipiiiciil and Ice linicpics as to make the completion of tests bv it at best cxtrcmeh dilTicult. Nevertheless, the picture can not be complelc'd without such analyses. Greater attention iherefore. must be concentrated on this phase of the problem. There arc also otlicT c|Mc>li(in> which luivc' a mcuc^ or less direct bearing on the relation of grouse foods to licaitli. In the light of wlial is known, iiow does food aflect weight, reproduc- tion and resistance to disease'.'' Is food one of the causes of periodic fluctuations in grouse |)opulat ions'.'' Such indications of the answers as are available are hc-re presented, but the many gaps ihal remain point lo tiiis licld as a parliciijarlv fniilfnl one fm future research. FOOD AND ITS RELATION TO HEALTH 237 Chemical Composition of Some Grouse Foods The various elements, compounds and organic substances that form the structure of a plant or animal are the items that make up its nutritive character. To determine the amount of each present a standard procedure is followed. First fresh samples of the food item to be ana- lyzed are gathered. From it the moisture present is extracted by drying. Then the oven-dried sample is chemically analyzed to measure the quantities of proteins, fats, fiber and ash or min- eral matter. All these are expressed in terms of weight per 100 grams of the sample. The sum of these is subtracted from 100 and the remainder, designated as "nitrogen-free extract"', is used to represent the more digestible carbohydrates, in contrast with the crude fiber, which is large- ly indigestible carbohvdrates. Such an analysis may be in itself extremely accurate. Because of the work involved, how- ever, it is seldom completed in sufficient detail to present a full picture of food composition. TREMBLING ASPEN g CHERRY O HORNBEAM Q. APPLE *" STACHORN SUMACH MOUNTAIN LAUREL EVERGREEN WOOD FERN BUDS ta~ ] n |i'::^- VJ-!-' r!.'-; ^^-.^""J^-:^i>,-,.- : ^- ,1 i-:'y. ■ , ., ■,: ., ~i LEAVES ■ I BUDS ■ 1 |C S ; CATKINS ■ 1 BUDS ■■ 3;H,J , o. -4 l_ , 1 BUDS ■ 1 f:>./ . • ■ : '-' ■ .V^ 1 ■ *- CAlKINsH j ; — -'; BUDS ^^M 1 l>-:.:v-- ■ ■■ ■ "'1 FRUITS ■ 1 tv.;. LEAVES ■ 1 :-'■:.' ^ '." ' ■•--^ "1 T ' ' 1 FHUNUS ^H 1 M...:X:.::...:.... ' ■ ^ -;..;V:.i | 30 40 50 60 70 PERCENT OF COMPOSITION MINERAL MATTER □ CRUDE PROTEIN □CRUDE FAT NITROGEN -FREE EXTRACT □CRUDE FIBER FIGURE 19. COMPOSITION OF EIGHT SPRING FOODS IMPORTANT TO GROUSE Nevertheless, it serves as a rough measure for establishing the amounts of these substances present in each food. Accordingly, the chemical composition of a number of grouse foods have been analyzed by the Investigation. The results are presented in lal>le 172 in the Appen- dix. For the sake of completeness and comparison additional examinations selected from published records but more or less applicable to New York conditions are also included. The proportions of these constituents present in eight foods commonly eaten by grouse in spring are shown in figure 19. One of these constituents, moisture, usually forms a larger part of a plant's structure than does any other item. It may be defined as the amount of free water not incorporated in other compounds. Its relation to health is discussed later in some details. Crude Protein This is the chemical compound most concerned with body building and feather develop- ment. However, proteins differ markedly in their ability to meet the varied wants of the body due to their fundamental differences in molecular structure. Dr. L. C. Norris of Cornell Uni- versity, working with the Investigation from 1932 to 1935. found indications that grouse chicks may thrive best on a starting diet containing from 27 to 30 per cent protein. This 238 FOOD HABITS AND REQUIREMENTS is considerably more than the 21 per cent level considered satisfactory for domestic chicks, but compares favorably with the 27 and 30 per cent that Psorris found were best suited to the needs of young pheasant and quail respectively. The comparatively large protein requirements of grouse chicks decreases materially with age. Those reared at the Research Center are now fed a ration containing 30 per cent pro- tein, which is dropped to 20 per cent for adults*. It is suggested that the need for protein by chicks parallels closely the proportion of insects found in the diet of wild birds. It has already been noted that insects make up about 70 per cent of the total bulk of chick foods for the first two weeks following hatching. This ten- dency rapidly falls off until by the end of August the food of young and adult are alike. Most insects are relatively high in protein content. Carpenter ants, one of the most com- monly eaten species, contain 30 per cent. Adults and older juveniles depend largely upon plant foods for their sustenance. .\s regards protein content, the leaves of aspens, dandelions and clovers, and the buds of cherries may stand them in good stead, for all are high in this respect as compared with most other grouse food plants, though low by comparison with most insects. It should be remembered, however, that digestibility varies with the kind of protein involved. As pre- viously explained, biological assays are necessary to determine the ability of the grouse to utilize the protein present in any given food. Fats These are considered so valuable a source of food that in dplcrniiniiig the nutritive ratio they are gi\en a value two and a quarter time.~ that of carbohydrates. The crude fat, or ether extract, given in chemical analyses represents the more or less digestible fats and oils as well as the less easily broken down resins and waxes, chlorophyll and other substances soluble in ether. Not all of the material here represented can be used by grouse. The crude fat content of the ration fed to the birds at the Research Center is 5 per cent for juveniles and slightlv less for adults. Even allowing for the indigestible fraction, a number of grouse foods contain well above the ration standard. The buds of huj)-hornbeam stand out in this respect as do leaves of mountain laurel. Surprisingly enough, hemlock needles and twigs, usually con- sidered of little value as food, contain a large quantity of f;U. allhough niiuh of it is probably in the form of resin. Sumach fruits are also high in fat content. Nitrogen-free Extract The term nitrogen-free extract is used to designate the more digestible carliohvdrates — sugars, starches and other saccharides. Since these figures represent the remainder from the other analyses rather than a separate analysis in themselves, small i|uuiililics of noii-car- bohydrates, such as tannin, are unavoidably included. It is significant that the foods most commonly eaten by grouse are all wdlsiipplied with these fat-producing and energy-releasing substances. The artificial ration led lo \ounu; birds contains 40 per cent, which is raised to 53 per cent for the adults. Spring foods, with the exception of hop-horn!)eam buds, together with all of the summer foods thus far analyzed compare favorably with this standard. The high proportion of such carbohydrates in the * See Chapter XI. |>. 489 fur feeding schedules. FOOD AND ITS RELATION TO HEALTH 239 spring diet is shown in figure 19. Unexpected is the high yield of the sumach fruits, con- sidered by many to be "starvation food". The leaves of some wood ferns contain even higher amounts. Crude fiber Under this heading are placed the indigestible carbohydrates, such as cellulose, lignin and chilin, as well as the resins and tars that are not soluble in ether. As such they probably are the least useful items in the picture. Unfortunately it is not known to just what extent grouse can digest this material. Most mammals can obtain little of value as food from it. although ruminents, such as cattle and deer, are able to extract therefrom some nourishment with the aid of certain bacteria. As might be expected, such winter and spring foods as buds, twigs and evergreen leaves all contain large quantities of crude fiber. A notable exception to this is the buds of cherry, which interestingly enough are taken more extensively in New York than is any other win- ter food. Apple buds and sumach fruits are also fairly low in fiber content. Here is a problem much in need of further study. Ash Under the heading ash is included the total measurable mineral matter that these plants contain. Since animal life is ultimately dependent upon vegetation for the majority of the inorganic portion of its make-up, the ash constituents, which provide the calcium, phos- phorus, manganese and other body needs, become highly important. Buds of apples, blades of certain grasses and fronds of wood ferns, all commonly taken spring foods, are excep- tionally high in mineral content. This is in fact characteristic of most grouse foods taken in late winter and early spring. The need for two of these minerals is indicated in that the cal- cium requirement for domestic chicks is 0.7 per cent; that of phosphorus 0.5 per cent; for laying hens it is 2 per cent and 0.8 per cent, respectively. Assuming grouse needs to be some- what comparable, those requirements are probably satisfactorily met at least in the spring diet. An element less prominent but exlremely important is manganese. Norris' experiments showed it to play a large part in the prevention of perosis or slipped tendon^"". This was formerly a connnon ailment of game birds raised in captivity, as well as in domestic fowl. It is described in somewhat greater detail in the chapter on Parasitism and Disease. The character of the soil strongly influences the mineral content of a plant. Wide varia- tions in the ash constituents of a species in different portions of its range are therefore to be expected. Aspen buds gathered near Ithaca, N. Y., contained half again as much mineral matters as did those analyzed by Hellmers'*", taken a little over a hundred miles south, in Pennsylvania. Vitamins In any discussion of nutrition the role of vitamins must play an important part. The artificial rations fed to captivity-raised grouse at the Center are, of course, fortified to insure the presence of these elusive substances in sufficient quantity, but their status in the normal diet is at present little understood. Nor is there much known of the basic amounts required for proper groulh and reproduction. The curing of a paralysis in grouse on experimental feeds by the use of vitamin B extract may mean that the chicks have an unusually high vita- min B requirement. Studies by Norris indicated the need of young pheasants for vitamin D 240 FOOD HABITS AND REQUIREMENTS and for tertain factors of the \ itarain B loinplex to be greater than that required hy domestic fowl. Their vitamin G requirement, likewise, was found to be from two to two and one- half times greater than that of white leghorn chicks. That such is the case for grouse does not necessarily follow, but a possibility is here indicated that awaits further investigation. Such is the general picture derived from the chemical analyses of a food item. Differences in Chemical Composition There are other equally important questions that might well be raised. One of them con- cerns the individual variation in these chemical components within the plant species that supply the food. As has already been pointed out. the character of the soil on which they grow plays an important part in determining the inorganic content of a plant. 2 TOP OF TREE *" NORTH SIDE U. BOTTOM ^ SOUTH SIDE U) BOTTOM t YOUNG SPROUTS it IN THE SUN WMmmmmmmimmMM IJaisJisiiiSiSJiMiSJi^iP" ^E mmmmMm,m. 30 40 50 60 70 PERCENT OF COMPOSITION MINERAL [—1 CRUDE I 1 MATTER L_J PROTEIN I 1 CRUDE FAT NITROGEN-FREE EXTRACT □ ?' CRUDE IBER FIGURE 20. DIFFERENCES IN COMPOSITION OF YELLOW BIRCH BIDS WITH RESPECT TO THE AGE OF THE TREE AND THE P.ART FROM WHICH THEY WERE COLLECTED Likewise there are substantial differences in composition between buds gathered from the shady as <()mpared with the sunlit parts of the tree. Results of the analyses of yellow birch buds taken from the top. the bottom on the north and on the south sides (all on the same tree) and from young sprouts of the same species gathered close by, are shown in figure 20. These examinations were made from material collected on April 6. 1942. Crousf are in great need of nutritive constituents in their food that produce heat and energy (luring the colder parts of the year. It is fascinating, then, to tind that the highly digestible carbohydrates that make up the nitrogen-free extract are somewhat greater in buds collected from sunn\ than from sha(l\ locations. Investigations by Gauman' " ha\e shown conclusively that this (ondition prevails throughout the year in tlic KurojH-an beech I Fapii.s sylvatica). Twig tips and terminal buds contain higher amounts of carbohydrates than do the branches and axial buds nearer the trunk. Those on the top of the tree ha\e the highest concentration (pf all. Here may be a likeK reason win grouse and deer feed on the buds from the top of ihc tn-i- when a\ailablc rather than fmrn the sliadcd >ide branches. F'ast growing sprouts, too, may differ from their slower growing companions as indicaled in table 172. Great variations are also apparent in the amount of crude protein, the buds from the bottom of the tree on both sides containing considerably more than those fnmi the top or fniin \oung sprouts. Seasonal differences, loo. have been noted. All such factors must, of (durse. be considered, in interpreting the residts of chemical anaKses of grouse foods. This. then, is a broad outline of the pirliin- that i> jii.-t beginning to take shajie. A large number of grouse foods still remain to be analyzed. The significance of changes in plant FOOD AND ITS RELATION TO HEALTH 241 composition should be determined. To answer some of the problems a far more complete analyses than the '"crude" determination of basic nutrients is required. The present studies must be continued for a long time to come before many relationships can be even reasonably well understood. Finally, the limitations of the data provided by chemical anlyses alone must not be overlooked. Knowledge of the nutritive constituents is largely academic until the abil- ity of grouse to use them is determined by biological assays. Relation of Food to Weight One of the less exact yardsticks bv which the relation of the diet to the health of grouse may be measured is the ability of a food to sustain weight and condition. That the birds are able to survive from year to year on their normal diet would seem in itself indirect proof of its adequacy. But. as has been indicated, such an assumption does not present the entire picture. The physiological experiments reported elsewhere* indicate that grouse have a re- markably high rate of metabolism. The amount of i-iicrgN they expend, even in a simple flight, is tremendous. Even the varied diet enjoxed l)\ grouse in .New \ork is not always sufficient to meet completely the normal demands of the bod\ when the energy output is high. Weight records of collected specimens bear this out. I he females show an almost con- stant weight loss from January until the end of the l)rccding season in June, and the males until after the moult. When these periods are completed, the liirds then gain steadil\. build- ing up reserves against the advent of cold weatlict. until ;i peak is reached at the beginning of the following winter. With the coming of snow and low tem|jeratures greater energy is required to move about and to combat the loss of heat from the body. Then, too, the birds are forced oflF the ground to feed in the trees. It is indeed fortunate that they arc then capable of existing upon browse, for an enforced migration in search of food might well prove disastrous. Yet there is no indication that the diet is wholly inadequate, for in no case was the weight recorded in a healthy bird dangerousK low. nor has the Investigation ever found a grouse in its natural habitat that died from star\atioii. Ihe normal weight-health relationship of New York grouse, adults and birds of the \ear. is indicated in tigure II. p. 90. It is interesting to find that the weights of captive birds at the Research Center likewise drop during the winter, despite the fad that the\ rccei\c an artificial ration calculated to best suit their needs. Noteworthy, too. is the fact that in winter and spring the quantities of food eaten by wild grouse is considerably greater than is taken in sunnner. Relation of Food to Cycles The possibility has been suggested that food, through cyclic qualitatixe changes in its nu- tritive components, may have a bearing upon the periodic fluctuations in grouse populations. It has been clearlv established that the composition of plants, on which the grouse must de- pend for food, may vary somewhat with weather conditions. Criddle' has reported on fluctu- ations in the abundance of grouse and other species of wildlife in Canada. DeLury"" has connected these observed variations with a close correlation between sunspot cvdes and some weather phenomena. All this is interesting but not even circumstantialh conclusive for. as indicated in Chapter VI, weather may have a host of other direct as well as indirect effects on grouse abundance. The weather changes most likelv to influence grouse food composition are precipitation, teni- * See AjipeiHlix. p. 719. 242 FOOD HABITS AND REQUIREMENTS perature, and the intensity, duration and composition of sunlight. Weather records of local stations, however, fail to exhibit such a cyclic trend in most of these factors. In New York, wide differences in the amount of precipitation as between individual sections of the State over the same year normally occur. No substantial rela- tionship was found when an attempt was made to link precipitation with subsequent grouse abundance from State records covering the past 40 years. Several records of high populations of grouse produced during abnormally wet as well as in dry years were found. But no studies of food composition on wet vs. dry years were made. Temperature also affects plant composition, but the extremes are more likely to be felt than the average. Many plants are capable of growing under wide variations, therefore a trend differing only a few degrees from year to year in the average temperature would be unlikely to have an appreciable influence. The most noticeable effect of temperature, to date, is connected directly with grouse mortality rather than indirectly through food. The third factor to consider is the relation of the intensity, duration and composition of sunlight to food. The first of these, changes in the intensity of sunlight, is closely tied up with sunspot activity and is, therefore, cyclic in nature. The possibility that light intensity affects plant composition, however, is largely offset by the phenomenon known as solarization. This arrests the action of chlorophyll as the intensity approaches a certain degree that is reached on all but the cloudiest of days. Duration of sunlight, on the other hand, although extremely important to plant growth, bears no relation to sunspot activity, for the length of daylight is normally much the same year by year. Sunspots control or affect still another physical condition, that of the quality of sunlight. Little is known of their effect on plant composition, though changes in the amounts of ultra- violet and infra-red light have been observed. The presence of vitamin D is dependent upon the former, but it is doubtful if quantitative changes in plant foods, thus brought about, are great enough to seriously affect the health of grouse. Experiments with poultry" have shown that a relatively small quantity of sunlight will offset the absence of this vitamin from the diet. It is noteworthy that the Investigation has never found a grouse, either in the wild or in captivity, suffering from rickets. In considering the broad picture it should not be forgotten that 994 separate food items have been identified from New York grouse alone. Then, too, their food is the product of at least two growing seasons. Buds and leaves are formed from the energy of past years. whereas fruits are more largely the result of the present year's activity. Such considerations tend to discount the probable effects of cyclic changes in ])lant composition on grouse abun- dance, even if such do occur. The diet, too, can influence population by afft'cling directly their resistance to the various decimating influences in their environment. A grouse in pot)r coiKlilioii is more susceptible to disease or ])r('dators than one in good health. It is possible that such a condition could be brought ai)out by a generally inadequate diet. However, in view of the foregoing discus- sion, no cyclic tendency in grouse abundance, due to diet, is indicated. Likewise dietary deficiencies could disrupt reproduction. Here again, there is no evidence to verifv a cvclic relationship. It is noteworthy that the number of eggs laid by New York grouse remained almost constant over the 13 years of the Investigation*. • See Chaplrr VUI. p. 360. WATER REQUIREMENTS 243 All this points to the conclusion that weather cycles are not likely to exert a strong indi- rect influence on grouse abundance by producing periodical changes in the composition of their food. But the subject is still too little understood to justify aught save an open mind as to the possibilities. WATER REQUIREMENTS Water is of prime importance to the living body, for it is vital to every physiological action as well as being an important constituent of the individual cells. Grouse may obtain water from three main sources: drinking water (from puddles, streams and ponds, and also snow), dew, and succulence (water taken from plant or animal foods high in moisture content). The extent to which the two latter sources can supplement or take the place of the former determines the dependence of the birds upon a steady supply of drinking water. The basic requirements of the species for water are not as yet definitely established. Ixmg. in carrying out physiological experiments*, however, found that a deficiency soon made it- self felt. Six adults were held without access to either food or water. After a period of only 3I/2 days half of them had died. On the other hand, 12 birds supplied with water, but no food, were able lo mniiilaiTi normal body trm|icraliirrs for nine days and easily survived the experiment. Birds reared at the Research Center are provided with drinking water from the time they are hatched. Inasnuich as their artificial ration is low in moisture content it is not surpris- ing to find them drinking freely. When the rearing of grouse was still in the experimental stage the chicks were fed moist foods such as (laljbcr and fly larvae. Drinking water, though available, was then onh occasionally resorted to. Adults penned in large enclosures also man- aged without frequent recourse to drinking water, dew and succulence apparently satisfying their needs. Small puddles formed by rain may be utilized by wild grouse. Bradbury'' witnessed such use but despite thousands of observations the Investigation has never actually seen a wild adult grouse drinking from open water. Lehmann™ similarly reported that prairie chickens rarely drink from surface water, but Girard''' states that sage grouse drink from two to three times daily, preferring running water to that from puddles. The sources from which water is available, may vary with the different seasons. In the spring melting snows and heavy rains make surface water everywhere abundant. A little later the succulent foods, to which the birds turn their attention after a winter-long diet of browse, are high in moisture content. Aspen leaves and fern fronds are well over two-thirds water. In contrast, buds and catkins generally contain less than 50 per cent moisture. The amount present in 8 common spring foods all freshly gathered, is shown in figure 21. Summer foods are made up largely of fruits and insects for the juveniles, fruits and leaves for the adults. All are high in moisture content. Raspberries and blackberries, the most com- monly eaten summer food, are four-fifths water. Dandelion leaves run as high as 89 per cent. The moisture content of many common fall foods such as beechnuts, hard seeds and buds is somewhat less. For example, sumach fruits, taken commonly at this period, are less than 10 * Sep Appendix, p. 761. A Bradbury, H. M.. persona! letter to the authors. 244 FOOD HABITS AND REQUIREMENTS per cent water. Roth in suimiipr arifl in fall the birds depend to some extent upon dew to help meet the hod) needs. Snow may take the place of dew to supplement the winter diet, which is also reduced in water content. Aspen 1 Yellow Birch Buds Leaves Buds Catkins CO ■^ Cherry Buds "-^ Hop- Buli ami open licl(l>. On the olliei- liand King's area in Miimesota. and the Adirondack unit arc rnad<' up of more or less continuously forested land. Many of the brushy areas here occur along the streams or in the swamps, .^ince these are the type of cover most utilized by New York grouse during the summer period, it seems possible that their distri- bution at least in the Northeast is not affected b\ water supply as much as by the associa- tion of |)lants that are often found in moist areas. There seems to be little difference between broods and adults in this respect. Likewise dur- WATER REQUIREMENTS 245 ing the winter period there was no substantial correlation between the distribution of adults and streams or swamps even on the Adirondack area. 9 \_oco*>on» m Drv V«uit of the American Part- ridge (Quail I. their dog |)ut up a Hulled Grouse. Both jicnileinen fired at the same in- stant— and seeing the biril lodge in a tree, both claimed tlie ]>ri/i' each supposing that his shot had taken effect. On arriving at the spot, they fouinl tlic liird impaled on a small dr\ branch, without having received a sinjrie ])cllet." Migration Observations during the present Investigation have revealed nothing to indicate that the ruffed grouse is other than thoroughly sedentary in its habits. NV'hile young birds may wan- * See Chapter \l, p. SO. COMMON TO BOTH SEXES 255 der considerably prior to their first breeding season, adults which have established themselves, seldom move far. Once settled, both their daily and seasonal ranges are usually small*, al- though in some of the more mountainous districts, particularly in the West, there has been reported'*'' "* a movement of grouse from the higher elevations to lowland areas in the fall and vice versa in the spring. Most references to migration in this species, however, either pertain to the so-called "crazy flight" or, more often, are erroneous interpretations of the occurrence of periodic scarcity. On the other hand, certain records of some of the older naturalists are worth citing. Audu- bon'' attributes to them habits akin to migration: "The Ruffed Grouse performs partial sorties at the approach of autumn", when they may be observed crossing such rivers as the Ohio and Susquehanna, "in parties of eight or ten, now and then of twelve or fifteen". He refers particularly to such movements from Ohio, Illinois and Indiana into Kentucky. He further states that, with the approach of sjiring, "the males, as if leading the way, proceed singly to- wards the country from which they had retreated. The females follow in small parties of three or four". While he speaks as though this were not an uncommon occurrence, subsequent observers have not noted it. Whatever the actual facts may have been, therefore, this trait is not now important. Other observers have, now and then, reported the occurrence of flocks or packs of grouse in the fall which seemed to be traveling as a group from one locality to another. For instance, Brewster^" relates finding a flock of one hundred and fifty to two hundred, in September, 1870. at the Forks of the Kennebec in northern Maine. On another occasion, he followed a flock of about 15 grouse over a mile through open farming country near Lake Umbagog, (Me.) be- fore he lost track of it still going in the same direction. More recently, Spiller"" writes of three similar observations, one of which involved a pack of 40-50 individuals which he wit- nessed personally. Having encountered nothing of this nature during the present study, it is difficult to inter- pret these experiences. All have occurred in the fall. Audubon suggested a quest for food as a motivating factor; Brewster a vestigal tendency to migrate. Spiller reported an otherwise general scarcity of birds at the time of his experience. Without more definite and better correlated data, however, one might speculate endlessly on these and other equally intrigu- ing possibilities. In any case, such group movements are rare. That local conditions may sometimes be responsible for apparentlv migratory movements of grouse is illustrated by the following account from Smyth"'" with respect to the vicinity of Ithaca, N. Y., in the fall of 1924: "Early in the fall, mast was very abundant and the grouse were widely scattered through the woods, appearing to be relatively scarce. Toward the close of the hunting season, however, hunters reported the birds as having become abundant, this being due to the gathering of the grouse to more favored feeding places as the picking from the woods in general became poor." Fall Dispersal (Crazy Flight) The so-called "crazy flight" among grouse has long been the subject of much discussion. In the fall of the year, birds are often picked up which have flown blindly against buildings or through windows, frequently killing themselves. Others are found in unusual places far from their customary haunts, as in the business districts of large cities or about houses and barn- yards. Their mad recklessness at such times may be judged from the fact that thev have been * See discussion under Territory, p. 258. 256 GENERAL HABITS known to crash through plate glass windows a quarter-inch thick. Forbush'" states that they have even flown into locomotive headlights. The impulse underlying this behavior is not clearly understood. Several theories have been advanced, at different times, to explain it. An anonymous writer in the Rural New Yorker, in 1875, attributed to an "old hunter" the opinion that the birds were frightened by the falling leaves. A writer in Forest and Stream, in 1878'"'^ states it is a provision of nature against detrimental inbreeding. Forbush"" suggests the possibility of an inherited instinct of migra- tion occurring sporadically among the birds. The same author credits Seton with the obser- vation that "'it is a trait of the young birds, which they exhibit during the first season and sometimes in the second, but never afterward". On the other hand. Gross'"" and Allen", find- ing infestations of the stomach worm (Dispharynx) in grouse picked up dead under similar circumstances, suggested the possibility of disease as the cause. While definite experimental data are lacking, it seems probable that this phenomenon is largely a characteristic of young birds seeking a territory in which to "paddle their own canoe". Of 15 specimens with this history which have been examined during this Investi- gation*, all were birds of the year. The time of usual occurrence in the fall coincides with the period immediately following the breaking up of most of the broods. Apparently, it repre- sents Nature's way of dispersing the surplus among the current season's crop. During this period, some stimulus, we do not know what, spurs the young birds to leave the family group in which they have traveled all summer and to strike out for themselves. At this time they become noticeably wilder. In corroboration of this, it is significant that, each fall, in late September or early October, the hand-reared birds in wire pens at the Research Center almost overnight became markedly more nervous and jumpy. This condition often lasts until Thanksgiving time. While such environmental changes as the falling of the leaves may have some minor influence, it seems certain that the controlling factor is inherent in the bird itself. Although the records are too scattered to be indicative, the incidence of "crazy flight" probably tends to vary directly with the fall abundance of grouse. Furthermore, it has been demonstrated that populations in excess of the saturation point decrease through dispersion at this season. Of problematical significance is the record of a bird marked by tying a bell to its wing, which was found dead sometime later, having broken its neck flying into a tree. This sug- gests the possibility that irritation resulting from injury or perhaps disease, may sometimes produce, in individual birds, reactions superficially resembling crazy flight. Spring Shuffle As noted above, the territory in which they pass their first breeding season becomes the permanent home of most grouse. The majority make this choice during the preceding fall. Some, however, either are laggards or become dissatisfied with the covert in which they spent the winter. The result is a "spring shuffle" in which such individuals wander about until they become settled. This niovcmenl. which takes place in late March and early April, is on a much smaller scale than thai of the fall and the birds seldom exhibit the wild recklessness of the crazy flight season. During the spring, birds are often picked up dead after having flown into some obstacle. Such instances have been thought, by many, to represent a recurrence of the crazy flight im- * Since the burta of Fabricut bai been employed a* an indel of age. COMMON TO BOTH SEXES 257 pulse. In most cases, however, such specimens have been found in or close to normal grouse coverts rather than being in some definitely "foreign" situation, such as a large city. Usually they have collided with a fence, telephone wires or even a tree trunk. It seems more logical, therefore, to consider them merely as accidents associated with the spring shuffle. Territory During recent years, students of bird behavior have, to a large degree, restricted the use of the term "territory" to denote that area which the males of certain species establish and defend just before and during the mating period"'' "". This concept is most evident among various migratory species whose breeding range is distinct from their winter range. But it may likewise be noted in non-migratory forms, although here it is often obscured by the fact that the same range is occupied the year around. On the other hand, especially among sed- entary species, the term may also be used to represent the area necessary to satisfy the habitat requirements of individual birds throughout the year and within which is included whatever specialized breeding territory the species may have. The former of these two concepts will be discussed first. Breeding Territory With respect to territory in this sense, Leopold^ suggests that "the ruffed grouse practices polygamy of Wight's 'crowing ground' type", in which the crowing ground of the cock tends to include the nesting territories of the hens which are more or less defended by him. Obser- vations in the case of the grouse, however, have failed to reveal any definite relationship be- tween the nest and the drumming log or evidence of defense by the male of the area in which the nest is located. This is in keeping with the fact that he has little to do with either the nesting female or the subsequent brood while, with the pheasant, there is considerable asso- ciation of this type. Rather, mating may take place anywhere within the cock's territory and it seems to be the responsibility of the female to seek out the male at such times, which results in such activity being most often associated with the vicinity of a drumming log*. While the latter may be defended, on occasion, against other males, the establishment of distinct breeding territories has been but slightly developed in this species. Year Around Territory In its broader sense, the territory of an adult grouse is that area within which the bird finds those environmental conditions it requires for a normal existence and within which it tends to remain permanently. As a non-migratory species, it seeks to satisfy those require- ments in a relatively small area. While other features are necessary, in varying degrees, it is obvious that the primary qualities which such an area must supply are adequate shelter and satisfactory food at all seasons and in suitable relation to one another. It is normal, then, for each grouse to establish its own territory. The areas occupied by individual birds, however, are extremely difficult to define. In the absence of conspicuously marked birds, the most significant records indicating territorial limits are those of birds which, when flushed, fly back past the observer. Unfortunately, however, instances of this type, in addition to being infrequent, most often occur at the edge of an open field, which is a recognized boundary anyway. For this reason, it has been necessary to rely mainly on successive re-flushes of the same bird and on repeated contacts with what is judged to be the same bird over a period of several weeks'^. Analy-sis of such plotted data demonstrates that, * See discussion under Mating Habits, p. 266. A See diacuesion of Methods and Techniques, p. 714. 258 GENERAL HABITS while individual birds probably have their own "stamping ground", the territories of several usiually overlap one another to some degree. This is especially true with respect to patches of conifers to which groups of birds resort for winter shelter. For this reason, any attempt to discuss the characteristics of grouse territory of this type must pertain to general areas used by several birds. Size. E^lsewhere in the book*, the requirements for suitable grouse habitats are discussed in detail. The size of areas (i.e. territories) within which groups of birds have been able to find conditions suitable for year around existence, has been observed to vary considerably. A major factor in this variability is the degree of interspersion of types. A separated unit, comprising 23 acres of diversified grouse cover on Connecticut Hill, has evidenced, through- out the period of the Investigation, a carrying capacity of one pair of birds and, in addition, has usually produced a brood. Although this unit is not sufficiently removed from nearby coverts to prevent some trading back and forth, it is the belief of the authors that, if com- pletely isolated, it could still support at least two grouse. From this, the size of these com- posite territories, as near as it has been possible to estimate them, has ranged through all gradations to areas of from 100 to 150 acres occupied by from three to 13 birds. It must be remembered, however, that a direct computation of the number of acres per bird over such an area rarely represents the size of the territory occupied by the individual members of the group. For example, although a population of 18 birds was estimated on some 90 acres of the Connecticut Hill study area in the spring of 1935, each bird, because of overlapping terri- tories, actuallv occupied a much greater area than five acres. The number of birds utilizing individual areas varies, in general, with the cover types present and their degree of interspersion. Of course, it also varies with the general population level. The figures noted above represent spring densities which are the best measure of car- rying capacity. In most cases, however, larger groups of birds were present during the early winter but became reduced through losses and through the spring shuffle. When Chosen. Final choice of territory by a grouse seems to be made i)rior to the bird's first breeding season. The locality chosen may be close to where the bird was reared or a considerable distance away. It is likely that this distance varies in general with the local population density. Similarly, it is probable that the so-called "crazy ffight" is a by-product of such movements^'. An interesting fact in this cuiiiiection is that adult birds, brought in and liberated on an area new to them, often "light out" anil travel considerable distances. But once settled, the territory usually becomes the peniianent home of the bird. This con- clusion is supported by an interesting series of data from female grouse trapped and banded at their nests on the Connecticut Hill area as illustrated in table 33. Aiiollier fciiialc. baiidid at her nest in 1932, was found dead in the spring of 1931 at a point 1,500 feet distant. Three other records of marked grouse have been recorded by I.l<)>d'". These birds were banded at Muscow, Saskatchewan, in March. 192.5. During the following October, they were shot at distances of a quarter-mile, a half-mile and a mile, respectively, from the banding station. The third case may represent either an exception or a first-year bird which had not become finally located when banded. • Sep Ch«|ilcr. 111. IV. una XV. A Sec dt*cu«tioD under Mobility, p. 25S. COMMON TO BOTH SEXES 259 TABLE 33. DISTANCE BETWEEN SUCCESSIVE NESTS OF INDIVIDUAL MARKED FEMALE GROUSE Year banded Year retaken Distance between nests 1931 1932 200 ft. 1931 1932 300 ft. 1931 1932 1.300 ft. 1931 1932 2,000 ft. 1932 1933 400 ft. 1936* 1937 660 ft. 1940 2.i0 ft. i93S 1939 1.150 ft. • Although this bird was not retaken in 1938 or 1939. the prea-nrc of broods from unltnown nesls in the immediate vicinity indicate she occupied the same territory. Relation to Cover Quality. It may be stated from the data at hand that tracts of as low as 25 acres of good cover* may afford a suitable annual range or territory for a pair of grouse but that, as the cover becomes poorer, the i)irds must travel over larger areas in order to find the enviroinnental requirements which they must have to survive. It is seldom, how- ever, that the territories of individual birds exceed 150 acres or an area, mughly. one-half mile in diameter. Therefore, the number of possible grouse territories in any tract varies with the quality of the cover — a relationship which is important in controlling carrying ca- pacity''. As might be expected, when a covert is not stocked to capacity, only the better territories in it tend to be occupied. For this reason, birds will be found in certain portions of an area, year after year, if they are to be found at all. while, elsewhere, one will encounter them only now and then except when high populations are present. Furthermore, the nature of individual units of this kind is constantly changing as the cover itself changes in the process of succession. Corresponding shifts in utilization by the birds also take place. Seasonal Shifts. Within grouse territories, there are seasonal shifts. In winter, the birds tend to concentrate in the vicinity of coniferous shelter while, in the sjiring. they spread out over the second-growth hardwood and overgrown brush areas. Likewise, in the late summer, the utilization of slashings increases and, in the fall, any portions of their domain offering fruits, such as hawthorns, wild apples and grapes, are favored. Gregariousness In 1812, Alexander Wilson'"" wrote "the manners of the pheasant [grouse] are solitary; they are seldom found in coveys of more than four or five together, and more usually in pairs or singly". While Wilson seems to have had little experience with grouse during our northern winters, his statement is very applicable to the habits of adults in the Northeast during most of the year. Although exceptions occur at all seasons, there is, in general, little evidence of sociability, except among the loose groups which occupy the same patches of shelter during the late fall and winter. Even these break up as the breeding season draws near. Illustrative of this situation is the record for the Connecticut Hill area comparing the fall and winter period of 1934-35 with the following spring. From October through March, a total of 1,678 grouse flushes were recorded, of which 379 or 22.6 per cent, comprising 76 contacts, * Sec Chapter III. p. 110. A See Chapter XII. p. 522. 260 GENERAL HABITS were found to be in groups of four or more. These groups were as follows: 36 of four birds each; 23 of five each; 8 of six; 4 of seven; 3 of eight and one each of nine and 11 birds. Dur- ing April and May, on the other hand, out of 1,327 birds, only one such group was encount- ered, this being of four birds in April. During the summer, it is very rare to find even two adults together. Several other observers, both before and since Wilson, have noted much greater degrees of gregariousness. Thus Morton in 1632,"" stated he saw 40 grouse in one tree. Forbush"" men- tions the report* of C.C.Abbot that, at one time, thousands congregated in the swamps of New Jersey. Again Lahontan^', in speaking of this bird to which he referred as "fool hen", says, "they sat in the trees in flocks and were killed one after another". More recently, Brewster™ and Spiller""'^ have reported large flocks in the fall. In view of the usual reactions of this species today, such records are striking by contrast. Undoubtedly, some degree of change in its habits has taken place, although it is very unlikely that large flocks were ever of more than infrequent occurrence. Furthermore, it is probable that many of the early records applied to broods. A common belief is stated by Billings"^. "The males form small parties and continue sepa- rated from the females until the approach of winter, when the males, females and young mingle together". The Investigation has secured no evidence of social groups of this kind among male birds at any season. Likewise, the supposition of Maynard™ that wintering groups represent broods which have not broken up, appears to be of rare occurrence. Rather, such groups merely represent birds whose territories include the same patch of winter shelter. Wariness One of the most important qualities which places the grouse of today in the first rank among game birds is its customary wariness wherever hunted to any extent. Primitive Tameness But this has not always been so. Descriptive of its original habits was the name of "fool hen" by which it. as well as its cousin the spruce grouse, was known in colonial days. The Indians hunted it with the club and with blunt arrows". Many early writers referred to it as stupid and stated it was considered fit game for small boys who most often secured the birds, not by means of firearms but by knocking them off their perches with sticks and stones, or by snaring them in a noose at the end of a pole. Another method is described by Nuttall"*: "They are even smoked to death, in the same man- ner as the wild pigeons in the western country, while sleeping harmlessly and unsuspectingly in their leafy roosts". Other accounts tell of groups of birds which allowed themselves to be shot, one after the other, without flying "■ "". • It teem* quite pnHsible that •nme confunion SB to nimes may have riiBtrd among Bome o( the early writing*. good auliiority that the heath hen onre occurred in the pine i>laini o( New Jersey in large packs, especially Abbot's report may in reality have applied to the latter species. A See discussion of Mobility, p. 2^^. It is known on n winter. Thus COMMON TO BOTH SEXES 261 The following account by Brewster", concerning the habits of ruffed grouse in Maine, is in- teresting : , "When I began to indulge in it [partridge shooting] about Lake Umbago in the early 70's the birds were exceedingly tame, even close to the settlements . . . My good dog found and pointed them readily, but was evidently not a little puzzled to comprehend why they should stand conspicuously upright in open ground, or on mossy logs, regarding him with seeming indifference from a distance of only a few yards . . . For instead of ris- ing promptly on wing as I wished, and expected them to do, they would . . . start off at a slow walk with crests erect and perhaps also widespread tails, shaking their heads and necks, and twitching their expanded ruffs at each deliberate step, and continuing increas- ingly to utter their derisive and unseemly snickering . . . Even when I forced them to take wing by running after them, they rarely went more than a few yards before alighting in a tree, or dropping again to the ground over which they might continue to hasten, if much alarmed, until it was useless to follow them farther." In this connection, Elliot^'" suggests that this extreme lameness is a characteristic of the Can- adian subspecies (B. u. togata). The Transition It is a far cry from the birds which prompted such accounts to the grouse with which most present-day hunters are familiar. But even today, in remote forest areas, one often meets with individuals which are comparatively unafraid. In the Adirondack* and other similarly forested regions, it is not unusual to find birds, either on the ground or perched in trees, which will allow approach to within ten yards or less. Similarly, one comes upon birds which merely run out of the way instead of flushing. Deer hunters frequently secure such birds with a rifle. It is also said that, in certain areas which have been set aside as sanctuaries, the birds lend to lose their wariness. But, in localities where hunting pressure has been appreciable, the species has undergone a process of education by which its members have acquired an instinct of caution. Beginning with the first pursuit of the birds by man, this process is gradual and continuous. Records do not tell us when the transition took place in most sections but, in his notes on the Lake Umbagog region of Maine, Brewster" states that, in that locality, it "began to be noticeable about 1889 or 1890", and that by 1900 the birds about the settlement had become "almost as wary and otherwise sophisticated as those found in eastern Massachusetts". Yet hunting cannot be held entirely responsible for this change. There seems little doubt but that the harassment of the birds by man's everyday activities is much greater in settled districts. Likewise, a similar relationship with respect to predators seems to exist in discon- nected coverts as compared with extensive forest areas. Present Caution In most coverts, today, these birds are thoroughly alert and wary. They usually rise well ahead and make off rapidly in a more or less zig-zag course which soon puts a tree or other obstruction between them and the observer. Again one will wait until it has almost been passed by and then flush in the opposite direction. Occasionally, they become confused and run directly at one before flushing. Like many other species of wildlife, they may often be approached quite closely in an automobile but flush immediately if one tries to get out of the car. After being flushed, they commonly alight in trees, particularly conifers. The female, while incubating or during the brood period, is, of course, an exception. When on the nest, she sits very close, probably depending on her protective coloration and immo- 262 GENERAL HABITS bility, but when with the brood, especially during the first few weeks following hatching, she is often remarkably bold and aggressive. Throughout the present Investigation, the majority of the grouse contacted have flushed within 50 feet of the observer. Over the fall, winter and spring period. 72.0 per cent of the flushes recorded have been in this zone, while during the summer, this proportion has increased to 87.4 per cent. Grouse generally flush much wilder in windv weather, however. Stratagems ^XTien trying to avoid detection, the grouse resorts to a variety of stratagems. The one practiced most commonly, perhaps, is to "freeze". If the intruder does not approach too close and continues on his way, the bird "sits tight"; otherwise, especially if one pauses in the near vicinity, it usually flushes. Sometimes, it will wait until passed by and then flush in the opposite direction, or again one will endeavor to sneak off by running. In hunting with a dog one frequently finds that a bird, upon becoming aware of the intruder, has run to the oppo- site side of some screening shrub or other obstacle before flushing. The apparent tendency for birds to dodge behind a tree or bush immediately after being jumped seems, in reality, to be most often accomplished in this way. An anonymous writer in Forest and Stream"" found one hiding in a hollow stump and a wounded bird was found by one of the authors to have taken refuge in a woodrhuck hole. On another occasion, J. V. Skiff. Deputy Commissioner of this Department, witnessed a grouse being pursued in the open by a goshawk. The bird flew directly into a thick pine tree successfully evading the hawk, but seemed to fall to the ground stunned. Yet a few moments later it was found unhurt. The ultimate seems to be represented by a wounded bird which submerged itself in shallow water until only its head remained above the surface. Forbush'*" has reported a similar rase. Another device is for a bird, roosting in a tree, usuallv close to the trunk, to draw itself up very straight and stiff with its neck upstretchcd and to remain motionless in this position. Brewster's" description of this trait is especially graphic: "Standing stiffly erect with raised crest*, grotesquely elongated necks and tightly-com- pressed body-plumage, they were easily mistaken for stout, stubby prongs where upward- growing limbs had broken off, or for thick clusters of Usnea moss, or perhaps for loose scales of rough bark." A particularly curious reaction, reported In a nuinbcr uf observers, is that of "treeing". Attributed entirely to the less wary birds of the backwoods, it is said that, when flushed by a dog, especially one which barks considerabh. they often merely fly into a tree and some- times remain there until the hunter can come up. Tame Grouse In contrast with the usual temperanicnl of thi> bird, the rintliiig. now and tiien. of a so-called tame grouse, gives a Dr. Jekyll and Mr. Hyde touch. One spring, as an observer drove up a little-used woodsroad near one of the Investigation's field stations, a grouse suddenlv emerged from the woods and ran beside the car. I'pon coming to a stop, it was fi>iiMd tiiat the bird would remain so long as the motor was kept running but flew when it was slnit off. This bird performed in nuich the same manner on several subsequent visits to the same spot. In another instance, a bird. a|)parently also attracted by the automobile motor, repeatediv ran alongside passing cars until caught one day in a trout landing net. Then, there was the case of "Billy and the Steel Mule"'. This bird was first encountered when it undertook to display, one spring, before a farm tractor. The latter, however, had to i)e running to be effective. There- COMMON TO BOTH SEXES 263 after, the performance was repeated many times and Billy often actually allowed himself to be picked up and taken for a ride. //. //. ( /r.ii. "billy" I see text) showing reaction to running motor of tractor •*i;:_3^;^*K;^:^ W. JT'. rrufii grouse attracted by wood chopping Nearly every year, reports are received of similar experiences. Some tell of birds frequent- in"; the vicinity of barnyards or farmhouses and. in some instances, allowing themselves to be 264 GENERAL HABITS fed by hand. One of the most common relates to the attraction of a wood-cutter's chopping for certain birds. One winter, while thinning a stand of voung cedars near Essex, N. Y.. Mr. J. R. Burnham noticed a grouse nearby which seemed completely undisturbed, either by his pres- ence or bv the sound of the chopping. Thereafter, for several weeks, this bird would appear each morning shortK after he had begun to chop and remain in the vicinity all dav, often hopping up on the log beside him while he ate lunch. And when he left at night, the bird would follow hini to the car and then fly alongside for a short distance down the road. A quite similar experience is related in the story of "Biddy" of Delaware County (N.Y.)\ After having made friends with a group of woodchoppers during the winter, this bird could be called throughout the spring and summer by simply "striking two sticks together" and, on one occasion, is said to have joined a family picnic in the woods and eaten a dish of rasp- berries**. Just what the psychological motivation of such reactions may be. one can only guess. Un- doubtedly, the sound of an automobile or tractor motor suggests the drumming of a rival but what associations are inspired by the staccato ring of an axe, remains a mystery. Feeding Habits The grouse is primarily a ground bird and its food throughout the greater portion of the year is largely obtained either from the forest floor itself or from the lower stratum of shrubs and bushes. Nevertheless, the birds are not averse to feeding at higher levels to secure the fruits of many trees and larger shrubs, when available. Furthermore, when budding during the winter, they commonly frequent the tops of large forest trees. So wide a variety of both vegetable and animal foods are taken that the species may be said to be to a considerable extent omnivorous.* Actions In general, grouse are constantly on the move while feeding although, when budding in the winter or feeding on fruits in the fall, they may remain for some time in one tree or shrub. In securing food from amongst the ground litter, these birds prefer to i)ick out what items they can as they move along without resorting to scratching as chickens and jiheasants com- monly do. In this connection, no evidence has been found to sujiptjrt the statement of Grin- nell'"° that "when the snow is not too deep, they scratch for food overlooked in autumn", or that of Forbush"" that the old birds are "persistent scratchers". When feeding on insects in the early summer along woods edges, trails and roads, or later on berries in the slashes and burns, they frequently junij) from the ground with neck outstretched and often aided by a stroke or two of the wings in order to secure items slightly out of easy reach. Another habit is to pick up ants which often abound in the material chosen for dust baths. An excellent account of their actions while budding in an ap|)le tree is given by Brewster", which is quoted in part : '■| ( iiiiiilid 11(1 less than nine m atlcicd all oxer the tree, a few being low down on stout limbs close to its main trunk and hciic-c inc(>iis|iicn(ius. but the greater number near the ends of its longer upper iiranches. where tlie\ could be jilairiK seen, while one or two were perched on the very topmost twigs. boKIK outliiu'd against the grey sky and looking as bic as hen-hawks. The\ were busily engaged in budding . . . At times, howi-yer. they UDiild all stand creel and motionless for a few monients. <'\idciitl\ looking and listening inlenth . Those feeding near the ends of long and slender branches had some difTiculty in • Src Cllninii l\ . |.. I'll. COMMON TO BOTH SEXES 265 keeping their foothold and were constantly obliged to jerk up their tails and flutter their wings in order to preserve their balance . . . They picked off and swallowed the buds in rapid succession, with much the same quick, bobbing motion of the head as that of a domestic fowl feasting on corn." Time of Day With respect to the periods of the day favored for feeding during different seasons, repre- sentative data are difficult to secure. It is traditional for the times preferred by game species to be early morning and late afternoon or early evening. With respect to the ruffed grouse, however, experience indicates that this concept is not wholly correct although, during the fall and winter, it may hold to a considerable degree. A comparison of the proportion of birds flushed at various times of the day in those cover types primarily offering food as against that in the primarily shelter types fails to show such a relationship during most of the year. This condition may be interpreted in one of two ways; either the birds have no regular feeding time or they find food and shelter sufficiently interspersed to render such a tabulation mean- ingless. While the former, undoubtedly, applies to some extent, particularlv in spring, sum- mer and early fall, the latter is probably the more significant. Similarly, observation of tracks in the snow indicates that in winter lhc\ often feed during the middle of the day. At this season, about as many birds are flushed at all hours from roosts as are flushed while feeding. During this period in particular, weather is an important factor, storms in general but wind especially causing grouse to restrict their feeding periods. On the other hand, there is considerable evidence to indicate that, when \isiting a certain s|)ot for the express purpose of feeding on some i)arlicular item available there, these birds utilize the early and late hours to a greater exlenl. This habit is particularly prevalent in the fall, a fact well-known to grouse hunters who turn it to their ad\antage. \gaiM. one winli-r. a group of four grouse was noted each evening, just before dusk, to fly into a certain large vellow birch to feed on the buds. This continued regularK for several weeks during February and March. Grinnell'" and Brewster'' record the same habit. Similar ol>ser\aliotis with respect to apple trees and thornapples have been noted. chiefl\ during the fall. Hut whether such obser- vatitms represent the whole of a given feeding period or merely one sto|) in ;i jicriod of much longer duration is not known. In an\ case il seems probable that this t\ pc of ri'lalimiship occurs ])ritnarily during fall and winter. Mating Habits In iSew \ ork the breeding season for ruffed grouse connnences in late March. Although drumming may be noted occasionally at any season, it is now heard frequently in all grouse coverts. Data indicate that the majority of nesting females lay their first egg sometime shortly after the middle of April. The period during which mating is at its height extends from mid-April through early May. although recurrences occur in the case of main birds whose first clutch is destroyed. 266 GENERAL HABITS In spite of the fact that a \(ihiminous amount has heen written relative to the courtship of the ruffed grouse, partirularlv with respect to the drumming of the male, few data have heen puhhshed coni'erning the actual relationships of the sexes in the wild during the mating period. Leopold** suggests that it is probably often promiscuous but stales that the usual situation seems to be a polygamy of a similar nature to the "crowing ground" t\ jie described by Wight"" in connection with pheasants, in which each male has his own separate group of hens. But the analogy holds only partially for, although the males establish drunnning areas "staked out" bv their drumming logs, there seems to be no recogniza])le relationship between these and the nesting sites of the females. The 484 nests, for which data are availalde from \9'M) to 1942, were located with respect to distance from the nearest kin lun druniniiiig log. ( See table 'M) . From this it seems that, if anything, the female selects a ncstiim site at a considerable TABLE 34. RELATION OF DRUMMIM; I.()(;S TO NESTS -CONNECTICUT HILL- 1930-1'>I2 Drumming Logs 0 to 50 foet 51 to 100 feet 101 to 200 feet 201 to 400 feet Over 400 feet Number 12 36 74 137 225 Per cent 2.5 7.4 15.3 28.3 46 5 distance from a drumming log. One year in the Catskills. however, a nest was but 10 feet from a used log. It appears most likely that, when in the mood for mating, the hen \ isils the cock in the vicinity of his drumming log rather than the cock going to the hen as in the case of phea- sants.* Supporting this conclusion, a number of obser\ations nia\ be cited. In a mimber of in- stances where drumming logs have been under observation, another bird (presumably female) has frequently been found roosting in a nearby tree, while the male occu'.ied his log. Once the male and another bird were found roosting together directly abovi' the log on several different nights and another time a second bird was observed on the (humming log with the male. On numerous other occasions, birds judged to be females have hec-n Hushed relatively close to drunnning logs while the male was drunnning. It does not follow, however, that actual courtship and mating necessarily take place close to the drumming log. There is no (piesticm but that more than one female often \ isils the same male, though no direct evidence has been secured as to the number which may nuUe with a single cock under natural conditions. On s<'veral occasions, however, two or more nests have been found inider circumstances strongK iri'■ -SWW^^T- COMMON TO BOTH SEXES 269 A badly vanquished bird may suffer severe wounds, chiefl) aboul the head and, unless separated, may continue to be the victim of attack until it finally succumbs. Instances have occurred, however, in which a bird died, not from physical injury but, apparently, from a form of nervous shock brought on by its lack lA ability to resist its stronger associates. In the wild, complete subjugation seldom occurs because there are always retreats to which a vanquished bird can fly and regain its composure. Moreover, the usual low densities of population provide a minimum of provocation for resorting to fighting. Aside from the breeding season, such combats take place most frequently among youngsters begimiing to "feel their oats" in the late summer and fall. During the breeding season, fighting most often involves males occupying adjacent drumming logs or in some way infringing on each other's "alleged rights". The males of this species do not indulge in contests for the posses- sion of the female as an individual. Instances in which fighting has been observed among wild grouse have been few and us- ually in(om|)lete. On one occasion, in October, one of the authors was silling quicth on a stump when two birds, one chasing the other, came running toward him. Tlie pursued bird soon flew, however, so no actual fighting took place. Again, earK one fall, Sumner M. Cowden, Superintendent of Fish Culture of this De|)artment. witnessed a clash involving three birds. two of which were engaged in a united effort against the other, which liriaily gave way and flew. This fracas lasted at least five minutes but aflcr tlic third bird Hi-w. the rcnuiining two seemed to be perfectly amicable. An unusual situation is reported by an anonymous writer in Forest and Stream"", who des- cribes a combat between two grouse on the same driiimniiig log in which the birds re|)ealedlv rushed at each other from a distance of a foot or two. After several "rounds", one bird retreated, jumped off the log and disai)i)cared. follow iiig which ibc ollii-r (Irummed once and then also walked off. Roosting While typically a nocturnal habit, grouse often roost during the da) time. In winter, espe- cially, they are fond of resorting to some protected spot where they can bask in the sun or sit out a storm. High wind, in particular, causes the birds to seek shelter at any time. In sum- mer, however, they roost to a much less degree during the day, although dusting may serve to satisfy any desire for a siesta. Grouse roost both on the ground and in trees, the greatest use of the latter being made in cold stormy weather. As discussed elsewhere* snow-roosting is commonlv indulged in when conditions are suitable but even during the winter they frequently spend (he night beneath a small conifer whose thick low-hanging branches form a sort of tepee. The shelter of a stumj) or log may also be utilized. For a tree roost, a thick-topped conifer is almost invari- ably chosen and birds have often been flushed from such perches as high as 50 feet from the ground. Their ability to take advantage of circumstances is indicated by Forbush"' who found several birds in a cave in the rocks, and Snnth" who reports a bird roosting beneath an old drain pipe. * See Hisnissiiin nf Snow Roosting. ] 2 0 GENERAL HABITS '"^^'0^^^, :^>^-.y,^k.%>^' PILE OF WliNTER DKOPPINCS SHOWING WHERE GROUSE HAD ROOSTED 0.\ THE GROLND FOR SEVERAL HOURS Although groups of several adults often roost in the same clump of cover, no tendency to form packs or coveys has been noted. While more than one bird may occu|)y the same tree, each is an independent individual. Snow Roosting Like the Eskimo in his igloo, the grouse may seek shelter during severe weather by burrow- ing into the snow. The impulse to do this seems largely correlated with marked drops in temperature or with high wind, although the presence of light, soft snow of sudicient depth is, of course, necessary. Forbush" also states that ground-roosting birds are at times completely covered by snow during a storm, thus creating the impression that they had burrowed into it. The habit has been observed throughout the range of the bird wherever heavy snowfall occurs. It is more ])revalent northward only because the necessary conditions are present more often. In New York, this habit is frequently employed at night during the winti r. Init the birds come out in the morning unless the weather is very severe. Individuals may. however, re- main in such a shelter several days, as evidenced by the number of driii)pings found in some burrows. During the (hntirric thev seldom resort to the practice unless the weather is very sharp and windy. When conditions are right, grou.-c ino.-t often di\c licmi an (i\criu'ail luancli oi. sometimes. even directly from (light. Such a plunge is made at an olili(|ii(' angle and carries the bird below the surface where a small space is hollowed out. often a foot or two from the entrance, if the while mantle is deep and soft. At other times, partiiularly if the snow depth is insuflicient to permit diving, the bird ma> walk In the chosen >\n>\ and burrow under the surface. Roosts of this type have been foiini! in snow only four inches deep, although this is unusual. The chamber formed |i\ ihc bird is slishtlv larger than its bodv and is seldom more than % '"^^^^T .T\nii,ii'^ ri) (liiANCiNr; Environment Fossil hunlcis lia\c found bones of the rufTed firouse among material of the Pleistocene age* from caves in Pennsylvania. California, Maryland and Tennessee, as well as remains probably referable to this species from Arkansas"". That it could survive such a period of time as well as occupv so extensive a range, both past and ])rescnt". indicates a high degree of ada|)ta- bililv. Certain further observations are. however, worth citing. Being a bird of the woods edges. • The rioiiitiit-riir ail*' "* B<*ul"Bic time rnilrd ;il...ul jri.OIHI vpor« ni;.t ami i* r-limiitril in luiv million year*. A Srr Cliaplrr II. p. 18. vfrcti n |M>rinil t»l a|i|iroxiiiiali-ly n COMMON TO BOTH SEXES 273 the opening up of the virgin forests b} early land-clearing operations, lunihering. and even fire when not overly severe, created conditions decidedly more favorable for its existence. When extensive areas of open land were produced, however, the grouse was. of course, ex- cluded. But where even small tracts of suitable cover were left, it continued to exist even within short distances of cities and the larger towns. Today it is found as a breeding bird within 15' '2 miles of New York City. Extensive tracts of cover are not required. As noted under the discussion of Territory, farm woodlots as small as 25 acres of suitably diversified cover will support grouse throughout the year. Any activity, however, such as grazing, which seriously reduces the amount of under- growth, tends to eliminate grouse whether the area is large or small. In the oil fields of southwestern New York and the adjoining portit)n of Pennsylvania, grouse are quite conmion over areas where wells, pumping stations, moving cables, and pipe lines are scattered throughout the woods and are contimiallv being tended by workmen. One year, a nest was discovered immediately under a cable line which had been in use several hours a day throughout the season and which jiroduced a loud creaking noise at each stroke. Perhaps the most significant recent evidence of adaplabililv is the maimer in which the species has reacted to hunting by man. A stupid "fool hen" where little sought after*, it has. over much of its range, developed a wariness which places it. today, among the most highlv prized game birds of the continent. TOLERANCK OF OtHER SpECIES Conflict between grouse and other native birds in New ^ ork Stale fcir the occu])alion of ;i given unit of environment, is negligible. In fact, it is very doubtful if such a condition ever existed to a significant degree. In anv event, the species, which at one lime might have offered competition, are now either absent or of very restricted distribution in the State. The wild turkey is no longer present, the spruce grouse has been reduced to the verge of extinction and the quail is confined largely to the extreme southeastern portion. (Competition with other native woodland birds has not been important. Neither, so far as it has been possible to determine, is there any indication that the natural- ized pheasant competes for the right to occupy individual coverts or for the necessities of life, such as food. Essentially, the habitats chosen bv the two species are distinct but. where swamps and woodlots are associated with the agricultural land use and mild climate required by the pheasant, both may often be found in the same coverts throughout the area where their ranges overlap. Nevertheless, no direct conflict between individuals has been noted. On the other hand, under such circumstances, the parasitic propensity of the pheasant of laying its eggs in grouse nests'^ may be an important reason for the scarcity of grouse in many coverts of seemingly high quality adjacent to lowland farms. Others have found that, where several species having similar habits do occupy the same habitat, there is a tendency for the limit to their combined increase to be reached at about the same level that any one might attain individually.""' Experience in 1931-32 and 1933-34, with the Reeve's pheasant (liberated in grouse coverts) corroborates this conclusion. * See discussion of Wariness, p. 260. A See Chapter VU. p. 334. 274 GENERAL HABITS CHARACTLRISTIC OF THE MALE Beyond the foregoing, there are certain important habits peculiar ti> llie male. They are primarily associated with Kreeding and that of dnimming is perhaps of greatest general in- terest. Drumming Wild bees liiini through the forest vines Where the bullets of England hunnned, And the partridge drums in the ringing pines Where the drunnners of England drunnned. Robert W. Cliiambers. No phase of grouse life has been the subject of as much description or occasioned more speculation than has the drumming of the male. Called the "carpenter bird" by certain Indian tribes'", the bird has since been aptly dubbed "the kettledrum of Nature's orchestra"^. In one of the earliest accounts of the habits of this species, Lahontan"'', referring to "their flapping", termed the performance "one of the greatest curiosities in the world". In a letter accompanying the first specimen of this species sent to England from Pennsylvania about 1750 by John Bartram (in Edwards'"') attention is called to the "remarkable manner" in which "they thump". The same author"' quotes a letter dated 1 7.52 from a Mr. Brooke of Maryland referring to the "beating of the Pheasant". Although Lahonlaii and others had likened the sound to that of a drum. Audubon"" seems lo have been the first to use this term directly to designate the performance. That this charac- terization was apjiropriate is attested by the fact that subsequent authors have used it almost without exception. With the rise in popularity of a number of sporting periodicals about the middle of llip niiieleenth century, notably Forest and Stream and American Field, discus- sions of grouse drunitning became mmierous and many an observer, both scientist and woods- Muiii, undertook to explain the manner in which it is produced.* 1 lie Druinininu. I'ri jitriiidnre In executing this curious invitation to love or war, the male grouse, usually from some slightly elevated perch, such as a log. stump or boulder, proceeds to beat the air with his wings, producing a hollow, ventriloquistic sound which commences as a series of distinct "thumps" followed by a pause, continues with the strokes coming "so close together that the sound of each merges with the next to produce the whirring of the 'nmffled drum' "^', and finally concludes with a single, weak beat. The cadence of the ordinary drum, expressed in sixteenths of a second between thumps, has been represented graphically by Allen ( in Bent™) as follows: .5-6-8-a-6-.'i-5-4-4-3-3-3-2-2-l-2-l-l-l-l-l-l-00()0()OnOO(K)()()()0()Oo6oO-l. Tin- time re- ([uircd has been delcrmitied In the same author as "almost cxactK eight seconds . Although members of Uic Investigation have observed and |ihotographed innumerable drummiiigs, it seems su|icr(liious to add another description of ihe performance to those already in the literature. Brewster'"', for example, has painted a particularly graphic picture of the preliminary stages as follows: "Suddcid\ he strclched up his neck . . . and glanced keenh around . . . Then he circled four or five limes around a space no more (ban a foot in diameter . . . and facing at a right angle with the log seated himself on it in the altitude of a Penguin ... As he sat thus . . . his tarsi and tail were pressed closelv for their entire li-ngth on . . . the log. A moment later the bird bejian to drum." • Scr llrrwMrr". niMlgc'"-. \lrrl«llil«. S.wypr^. Allpil (in llonl*') . CHARACTERISTIC OF THE MALE 275 Richard E. FolUu A DRUMMINr, GROUSE The balance of the performance is described by the same author'^ as follows: "a quick stroke [was] given forward; the wings immediately recovered and another stroke, a trifle quicker than the first, was succeeded by another still quicker, until the wings vibrated too fast to be fnlbiwcd by the eye . . ." Regarding the initial thuni|)s. Allen (in Rent™) says: "he appears to throw his 'shoulders' back. This might give the impression that the wings were struck behind the back, because the forward stroke of the wing follows so instantaneously that the eye scarcely perceives it. and it is given with such force and the wings come back to the normal ])iisition so quicklv that the entire action registers on onlv one frame of the motion-picture film having an exposure of appruximately one-fiftieth of a second." 276 GENERAL HABITS I he drumming perfonnanie unavoidably advertises the bird's whereabouts to enemies as well as to others. It is significant, therefore, that a bird can stop in mid-action at anv time except durino: the height of the "roll". Individuals have been seen to "freeze" in this manner at the approach of a dog or upon spotting a hawk overhead. Even the faint click of a well- muffled movie camera will cause this reaction. Nevertheless. man\ birds arc killed each vear on their drumming logs. The Sound Many times during the Investigation, field men have mistaken the sound of a distant gasoline engine or tractor for a drumming grouse. Over the vears. besides likening it to "the rapid rolling of a distant drum " ', a variety of other similes have been used by different observers. Thus, Lahontan"'' speaks of it as a "humming noise", and Edwards"", quoting Bartram. re- ported "they sound like thunder at a distance" — a comparison which has since been made by iimumerable authors. Its resemblance to the booming of a far-off gun has also been suggested. Ventriloquistic to a high degree, the sound is deceptive with respect to both the direction of the drummer and his distance away. Over the hilly, broken terrain of most grouse coverts in the Northeast, it is most often audible at distances up to somewhat over a quarter of a mile. Under especially favorable conditions, however, it has been heard nearly a mile awav. The direction of the sound tends to be more confusing to an observer within the woods than to one in the open. Apparentlv, it echoes to some extent among the tree trunks. How Produced. Impelled by a certain air of mystery surrounding the performance because of the wariness of the birds, observers have from the beginning attempted to explain the method by which the sound is produced. Certain Indian tribes believed the birds beat upon a log with their wings and thus called the species the "carpenter bird"'"". Lahontan "' thought they struck "one wing against the other". Bartram"" pictured them as "clapping their wings against their sides", a belief shared by Audubon"^ and \uttall"". Brooke"' suggested "they swell their breasts like a pouting pigeon, and beat with their wings", but left it ambiguous as to whether the wings beat upon the inflated breast or merely the air. An anonymous author in American Field'"" suggested that the wings beat u|)on the inflated lungs, while another writer*''" stated the sound is jiroduced h\ "inflating an air sac . . . striking it with their wings and forcing the air out through the mouth". In fact. Mavnard ' excn contended it to be primarih vocal and that "the wings merely aid in producing it . . . just as a rooster crows, flapping its wings at the same time". In 1874. however. Brewster'' |)ublished an account of a series of careful observations of a drumming grouse from a blind but 12 feet awa\ and advanced defi- nitely the conclu>iiui that the sound resulted from the concussion of the wing-beats on the air alone. So the controversy went. The "hollow log theory" was earU ilispro\ed 1>\ the fact that grouse frequently drum on boulders and similar objects. Neither was any appreciable degree of credence placed in the possibility that air sacs or vocal sounds were involved. On the other hand, while considerable belief persisted in the theories of Lahontan and Bartram, the obvious care with which Brewster's observations had been made and recorded gained wide acceptance for his explanation iitilil I'JO.S. In that \ear Hodge' threw additional fuel on the fire b\ introducing pholograpliic e\ idence which he contended showed the sound to be produced b\ "the impact of the stitlU held concave wing on the feather cushions of the sides". At the same lime. Jones''" joined Brewster in the lielii'f that the wings beat against nothing but air. ^rsr- 3 *.l.v I \ the powerful forward, upward and somewhat inward stroke of the wing followed by its instantaneous reversal of motion. The reaction is some- what akin to that which produces a clap of thunder or the crack of a whip. In creating a sufficient displacement of air, the concave nature of the underside of the wing, together with the angle at which it is held, are undoubtedly of great importance. Purpose An attribute of the male grouse only, drumming is fundamentally a corollary of courtship and functions to announce the whereabouts of vigorous cocks both to females and to other males. Called "tlic throbbing heart, as it were, of awakening spring'"", it is indulged in most commoidx' during the early breeding season. On the other hand, it has been heard every month of the year and often becomes quite frequent in the early fall. In the spring, it may serve either as an invitation to the female or as a challenge to the world in general. At other seasons, it undoubledly represents an expression of exuberatice on the |i;irt of an utiusually vigorous individual. The Drumming Season Over most of New York, the drumming season begins in late March and reaches its height during late April and early May. At the higher elevations in the northern part of the State, the kettledrums commence a week or so later. Observation of grouse in captivity at the Re- search Center reveals that they must learn to drum. Apparently they instinctively know the motions but, even after acquiring adult ])luinage. require considerable practice to develop the ability to drum with tin- normal sound effect. On several occasions young males, which never attempted to drum wliilc in a mixed grou|). have been placed in individual pens with the coming of spring. These birds have usually jiroceeded to give a perfect drumming per- formance excejjt that there was little sound. A biid reported by Allen" took two weeks to learn. Thus, it might Ix- that the staggered dates on which one first notices various wild birds commencing to drum may be due. in jjart. to tlie young birds in the population, although the comparative vigor of vaiioiis individuals is probaliK llic prirnaix reason. TTie predominant drumming periods during llie spring are earl\ morning and late after- noon. Beginning usuall\ ucll before daylight. iIk- performance is repeal<"d with remarkable regularity until sunrise or somewhat thereaflcr. Mlbongh at the height of the season inter- CHARACTERISTIC OF THE MALE 279 mittent activity may often be heard throughout the day, it commences with regularity again about an hour before twilight and continues until dark and sometimes later. The frequency of successive drums increases as the height of the season approaches. Grouse have been ob- served to drum regularly for several hours at from 3- to 5-minute intervals. Birds have also been recorded which drummed for periods of half an hour at intervals of less than a minute and on several occasions have been seen to drum a second time with hardly a pause between. V^-^'. tr. :m Ri^f''-rt U . Uarrnw WHERE A GROUSE DRUMMED IN THE SNOW ON A GRANITE LEDGE IN FEBRUARY During this Investigation, few instances of fall drumming approaching the regularity re- corded by Brewster™ have been observed. The Drumming Log For a bandstand, a drummer usually mounts a fallen log, hence the term "drumming log". Most commonly chosen is some old, more or less moss-covered relic of fairly large diameter mouldering on the forest floor. Sometimes an old stump or upturned root is used, but freshly-cut logs or stumps are seldom favored. In rocky country such as the Adirondacks, boulders and ledges frequently serve this purpose. Birds have also been observed to drum from most any small mound on the forest floor, even stones or bunches of moss no larger than the crown of one's hat. It is probable, however, that in such cases, the bird usually has a more characteristic rostrum elsewhere, since individual males usually have more than one drumming station. Perhaps the most unusual "log" is recorded by Grinneir% who found a bird drumming on the carcass of another bird which had been shot and unrecovered a few 280 GENERAL HABITS days before. In captivity. o;rouse experience no difliculty in druniniint; on the flat wire floors of their pens. Little preference as to the location of dniniminp logs seems to be shown, except that they are usually found in woodland cover and where possible associated with conifers. Open or overgrown land types and steep slojjes tend to be avoided. A TYPICAL GROUSE DRUMMING LOG During the drumming season, logs in regular use may be recognized b\ the presence of numerous fresh droppings on or beside them. If such droppings are in some way obliterated from a series of logs and the subsequent accumulation noted at the end of a definite interval, one may secure some indication of the degree of use of each. This varies with the mimber of logs per male, the amount i>f competition, size of territorv. and other factors. A curious incident was noted on the Adirondack area when a bird, whose favorite log was located in a swam|>. contimied to utilize it while it was complelciv surrounded bv water from an overflow- ing brook. Manv logs are used vear after vear. Number of Logs Used. Male grouse usually establish iiiorc than one dniiinning log. In ad- dition to individual differences between males, there is some indication that the miinbcr mav tend to \ar\ direclK with the di'grcc of competition with other males, and inverscK with the relative mimber of females influenced. On the olhci liaiid. in several instances, isolated pairs CHARACTERISTIC OF THE MALE 281 have had no known drumming log, although a brood later materialized in the territory. It cannot be assumed, however, that the males of such pairs did not drum, as indistinctive objects are often used for drumming and. if the performance occurred but a comparatively few times, it might go entirely unnoticed. Nevertheless, among hand-raised birds, males which are not separated from the opposite sex seldom drum, although they usually mate successfully. During the course of the Investigation on the Connecticut Hill area, 1.566 drumming logs have been noted. In the aggregate, 1,173 male grouse have been present during the same per- iod. Thus, an average of 1.33 recognized logs per male have been utilized during this period. At the same time, in a number of instances, ratios much lower than this have been observed for certain groups of males with no loss of breeding success among the females involved. For instance, in one covert on the Connecticut Hill area in 1932. a total of 13 males established but two drumming logs, yet nine of the 1 3 females were known to have nested. On another portion of the same area, in 1934, but one log was found in spite of the presence of three males, yet all five females nested. Similarly, in 1939, only one of six males established a log. Apparently then, mating may often be successfully completed without the aid of an estab- lished drumming log. although as noted above, this does not mean that such males did not drum. Conversely, other males have used as high as three and four, or even more, logs apiece. In such cases, the relative exuberance of individual males is undoubtedly a factor. Of the various logs used, however, not all are resorted to regularly. In 193.'5, a separate record was ke|)t on the Connecticut Hill area of the logs in regular use as against those used only occasionally. It was found that 15,S males used a total of 214 logs, but only 142 regu- larly or .92 per male. Thus, it appears that some males, each year, for one reason or another, do 111)1 drum regularly on a definite log as is the habit nf the majority. Courtship Courtship among birds comprises those actions, performances and displays, employed chieflv by the male (except in a few species such as the phalaropes). which serve to bring about the pairing of the sexes and ultimate mating. In the ruffed grouse, it consists primarily of strutting and feather display*. At the same time, largely similar if not identical performances, except for the head-twitch- ing stage'^, are employed throughout the ^ear as one means of maintaining a bird's "social rank"'' or merely to show off. A modified form of display has alsobeen observed in females attempting to dominate weaker individuals. There is. therefore, no clear-cut objective means by which to distinguish courtship activity although the feather display associated with it is usually more complete. Nevertheless, at this season, such exhibitions, together with drumming which is regarded as a corollary, serve to bring the sexes together and undoubtedly, to some extent, to convince the female of the merit of the male. Thus they may be considered to fulfil the function of court- ship even though essentially the same display may be used for other purposes as well. Specifi- cally, they serve this purpose when witnessed by a female in the proper physiological condition for mating. In other words, they supply the psychological elements needed to complement the physiological. They reach their fullest intensity in the innnediate presence of a female in "heat". * See discussinn of Strutting, p. 282. A See discussion n( Head -twitching, p. 283, t See Chapter II. p. 63. 282 GENERAL HABITS \^ hether or not a female, having witnessed such a performance, mates with that particular male, is immaterial. In no species does courtship guarantee acceptance. Then, too, one must be very cautious in crediting conscious forethought to the activities of any form of wildlife and remember that, among grouse, the male does not seek out an individual female to whom to address his attentions. Furthermore, in a polygamous species, the association of two indivi- duals for mating is short. To other males, or fgmales not in the proper stage of the reproduction cycle, actions of this kind merely represent a "chip on the shoulder" of the performer. W hat seems to be the nearest approach to actual observation of the courtship of a wild grouse is recorded by Allen^. In the course of a study of drumming, this investigator placed a captive female in a wire crate beside a drumming log in front of the position usually occu- pied by the male. Upon coming to the log, the male, without drumming, soon began his strutting antics, accompanied by a full plumage display. This was continued for three-quarters of an hour, both on the log and on the ground beside the crate, "when he attempted to mate with the female . . . though the wire separated them by several inches". Following this, the male proceeded to drum, apparently as a challenge to any other males which might be in the vicinity. Strutting Strutting and the feather display which accompanies it, has two major functions. First, it is employed throughout the year by individuals of both sexes for purposes of showing off or expressing exuberance and as one means of asserting their self-sufficicncx among their fel- lows*. Second, during the breeding season, it also serves as the courtship of the male''. In the former sense, the performance has been termed the "intimidation display" by Allen". When used to express exuberance or cockiness, the presence of other iMdi\ iduals. regardless of sex, appears immaterial. In its fullest development, as given before a receptive female in the breeding season, it con- sists of a display in which the male raises his tail almost perpendicularly, spreading it fan- wise, while at the same time lowering his wings until they drag. Though the crown feathers are not raised, the head is held back and the rufi about the neck is erected until it forms an almost complete circle of iridescent black or brownish feathers. The body position is not un- like that of the strutting turkey gobbler. With slow and measured steps he ap|)roaches the ob- ject of his attentions, occasionally lowering his head and stretching his neck to ])eck violently several times at the ground or other objects before him. A slight Iiiss oftcii accompanies the forward movement. As excitation increases, the male stands still, though boih hiss and move- ment of the head become more jironounccd. each action being clearlv distinguished from the one that follows by a pause. Tlien the head is tilted slightly forward and the neck rotated with increasing speed from side to side as though pivoted on bodv and beak. The sound now reminds one of a locomotive rather rapidly getting under way. Abruptly the motion ceases and. with a prolonged, loud hiss and a dragging of wings, the male completes the action with a short, determined rush at the female. Even before a receptive female, there is considerable variation between the actions of dif- ferent males and. under other circumstances, a wide range of modifications is employed. Es- peciallv if no female is evident, a male often stints \\\i ami ddun his drumming log, either • Scr Cliniil.r U. p. 65. A S«e dilcuMion of Courlvliip, p. 261. I CHARACTERISTIC OF THE MALE 283 before or after druniniing. with merely lowered wings and perhaps slightly raised and spread tail. Young birds, particularly in the fall of the year, bluster and show off in a similar man- ner, frequently accompanied by a short dash at another individual. Evidence of the same type Gardiner Bump A STRLTTIAO OKOLSE of reaction may be observed during the winter and particularly in early spring, where snow lingers on the ground, when tracks are often noted flanked on either side bv a narrow cut in the snow left by the trailing wings of the bird. The display of the female is usually much less vigorous than that of the male and seems to be used only for maintaining her social status. Experience with captive birds, however, has shown that the degree to which a male dis- plays is no criterion as to his quality as a breeder. Frequently, highly demonstrative individ- uals prove complete failures, while others which seldom display are often consistently effec- tive. Head-twitching As described elsewhere* the male usually becomes quite gentle during the final stage of courtship. Tail down, he follows the female slowly about, pausing when she pauses. At the * See Chapter II. p. 67. 284 GENERAL HABITS same time he frequently shakes his head and neck sliphtly from side to side. This aetion is in no wise violent as is that whiih accompanies strutting. Rather the head, which is held some- what back toward the shoulders, is twitched with a rotating motion as though it hung on an inclined pivot running through the base of the skull. Relation to Nest The Investigation has been unable to discover any relationship between the male and the nesting site of the female*. Birds judged to be male grouse have now and then been flushed near a nest but there is nothing to indicate that such occurrences are other than coincidental. Among birds held in captivity under natural cover conditions at the Research Center, how- ever, several males have exhibited a tendency to sit on the eggs. One individual aclualK in- cubated a clutch until it hatched and then undertook to brood the chicks. A similar incident was observed at a farm operated by the American Game Protective Association"', and a third has been reported by Allen'". Relation to Brood Similarly, it has been impossible to discover any consistent relationship between the male bird and the brood. Nearly every summer, however, there have been a few occasions when a bird judged to be a male has been flushed very near or even directly with a female and her chicks. Most of these observations have occurred during the first month following hatching and the circumstance has never been recorded twice for the same brood. In one instance, the second adult clucked and strutted about while the female attempted to lead the intruder awav. Several other observers have reported similar experiences^- ""• "". Loveland^" states the male even went so far as to fly at him. Again, after a brood has been flushed, it has sometimes been noted that an adult, other than the brood female, will fly in and attempt to silence chicks which commence to peep. Whether such individuals were male or female could not be determined. It seems doubtful if occurrences of this kind are other than accidental or possibly, in some cases, representative of unusual individual variation. On the other hand, the second bird may have been another female which had either lost her brood or merged it with that of the first. CHARACTKRISTIC OF THE FEMALE As with her mate, the female also indulges in a number of acli\itics clKuactcrislic oidy of her sex and they are. likewise, chieflv associated with the r<'produ. '2M). CHARACTERISTIC OF THE FEMALE 285 rectioii were also recorded. On the other hand, in 1938, it was several days earlier, resulting in a total difference between 1938 and 1939 of over a week. Although the physiological devel- opment of the bird at this season is fundamentally activated by the progressive, dailv increase in the amount of light, which is constant each year, the onset of nesting seems to be influenced, also, by annual variations in weather conditions and. possibly, the availability of certain foods. But while the majority of females in a locality commence egg laying within a few days of each other, a normal span of roughly two weeks between early and late individuals has been noted. In the advanced season of 1938, a number of clutches were recorded which must have been begun before April 15. Several similar nests were found in 1941 and 1942. Conversely, in 1932, many hens, even down state, did not start their first clutch until after May 1. In this connection, there seems to be some tendency for birds, destined to produce large clutches, to start laying ahead of the average. Beyond this, greater individual variations often occur. The earliest date* recorded by the Investigation and, curiously enough, in the Adirondacks of northern Warren County (N.Y.), was April 7. Since this nest was broken-up before being completed, the number of eggs which would have been laid remains a question. Next to this is a record* of A])ril 8. fnim Dutchess County (N.Y.), for a clutch of 13 eggs. In general, outside the higher Adirondacks, clutches begun after May 1 usualK represent renests. Outside New York, the nesting season ajjpcars to be somewhat earlier as one goes south. Thus, several reports from southern Petnisylvania indicate the usual hatching date to be about a week earlier than the June 1 average encountered by the hnestigation. In interior Alaska, however, Dice'" found a brood on June 8. And Preble"" found females with voung on June 15 at Smith Landing on the Mackenzie River. Structurk ok TiiK Nkst The ruffed grouse is a ground-nesting bird. The nest is a \er\ crude and unpretentious affair. It can hardly be called a structure, as it is merely a cup-shaped depression formed by the female to fit her body among the leaxes of the forest floor. There is no evidence that any of the components are ever transported to the nest site from a distance. It is lined and the edges built up of materials within reach as she sits in the depression. A few contour feathers are usually found in the nest, but it is not believed that they are placed there deliberately. Because hardwoods are the major component of the shelter utilized for nesting in New York, leaves of this group almost invariably form the nest. A number of instances have been noted, however, where evergreen needles predominated. Its forination is essentially complete before egg laying commences. With respect to the actual size of the nest itself, six typical examples on the Connecticut Hill area were measured. The dimensions of these averaged as follows: diameter, 6I/4 inches; depth, 2''^'4 inches; thickness (of the walls). '4 inch. The most frequent location for the nest is at the base of a tree or stump while sites under a log, at the edge of a brushpile or at the base of a boulder or rock ledge, are frequent. Characteristically a depression on the forest floor. Smyth"' reports one "that was raised about six inches from the ground, being built in a kind of little pocket formed at the base of a ten inch tree where several small sprouts grew out to one side". Similar situations have been encountered during the present Investigation. Others have been found in or on low decaying * Estimated un basis of number of eggs fiiuiid in nest wliile female was still laying. 286 GENERAL HABITS stumps. Someliines quite unusual sites are chosen. One nest was found inside a large stump, the interior of which had rotted out in such a way that entrance was possible through an old axe cut on the side while the top surface was still intact. Several have been fciund under large y / 5? ri^-i^;^-: THE NE.ST OF A RUKKKD GROIISK IS MERELY A CUP-SHAPED DEPRESSION AMONG THE LEAVES OF THE FOREST FLOOR pieces of bark which formed roofs over them. Nothing, however, has been observed to support the suggestion of Samuels™ that, if persistently molested, the grouse will utilize the abandoned nest of a crow. The immediate site of the nest is dften remarkably exposed, the i)ird a|)parently relying to a great extent on her protecti\c coloration. Some, however, are in rather thick situations among sprouts, bushes or brush, but. in such cases, usually have an easy avenue of escape. No instance has been recorded in which the same specific site has been used more than once. Habits During Laying and Incubation While laying her eggs, the female grouse spends a minimum amount of time at her nest. Observations, corroborated by records ])erlaining to captive birds at the Research Center, in- dicate that, on the average, egg-laying proceeds at the rate of two eggs every three days. Thus, approximately 17 days are consumed in laying an average clutch of 11 eggs. CHARACTERISTIC OF THE FEMALE 287 Early in the season, comparatively slight disturbances often result in desertion of a nest. On a number of occasions, birds have been flushed from nests before a single egg had been laid, and, in a few cases, have deserted, apparently as a result of the one contact. The same thing has also occurred with respect to partially complete clutches. But desertion is, in gen- eral, infrequent and the great majority of birds persist in their nesting attempt unless the clutch is actually destroyed.* Prior to the beginning of incubation, nesting females flush more easily and at a greater dis- tance from the observer than later, ahhough there is a tendency for birds which have been repeatedly disturbed to be wilder than other- in the same stage of the breeding cycle. As the incubation period progresses, she sits increasingly close until, toward the end, one may practically step on her before she will flush. A number of instances have come to our attention where birds have remained on their nests in spite of logging operations immediately about them. In other cases, road crews, work- GROUSE WHICH CONTINUED TO SIT ON NEST IN SPITE OF WOOD-CUTTING OPERATION DIRECTLY BESIDE IT ing within a few feet, have failed to disturb sitting grouse. Perhaps the greatest determina- tion was shown by a bird in Orange County which continued to sit during a forest fire and even allowed a spray of water to be played over her. Similar experiences are reported from * See diBcusBion under Desertion, p. 290. 288 GENERAL HABITS I'eiinsylvania"'. in one of whiili llie bird is said to have fought the ilames with her wings. A curious activity of the female during this season is related by Allen'", who found the ap- proach of nest building to be "first indicated by feeble attempts at tossing leaves over her shoulder onto her back as she sits or even as she walks". The same author continues as follows: "The leaf-throwing continues during the egg laying period and for several days after incubation starts, so that at first she sits with her back mure or less com[)letely covered. This doubtless account.-, fur the differences noted in the covering of the eggs with leaves amongst wild birds." No indications have been found that grouse inlt'ntionali\ (d\cr their eggs when leaving the nest. The popular belief that they do has apparently originated either from the habit noted above or because a bird flushed from her nest will, now and then, stir up the leaves so that some settle over it. The attachment of the female for her nest and her fearlessness in attempting to protect it and the subsequent brood from harm represent one of the best recognized habits of the species. Not only does she sit so close that it is occasionally possible to touch her before she will stir, but when finally flushed she often flies only a few feet, then tumbles to the ground and attempts to lure the intruder away by feigning severe injury, often accompanied by distressing whines. Pretending to have a broken wing, she half hobbles, half flutters, along the ground. If the in- truder is deceived and follows, she keeps just ahead until a distance of some fifteen to twenty yards has been covered when she suddenly regains her power of flight and departs. Otherwise, she usually remains in the vicinity, fussing and clucking, or she may even attack certain |)red- ators as in the lase of a black snake as reported in the PennsvKania Game iSews'"". This "broodiness", however, usually develops gradually. primariK during the latter part of the incubation period and does not become pronounced until a few days before the eggs hatch. But, in a few instances, a tendency in this direction has been noted early in the second week and, in one case, a slight attenijit to distract the observer was made by a bird which had still to lay the final eggs of a clutch of fourteen. Incubation in this species iiormalK requires a|)pr()ximately 21- days, varying between 23 and 24. a determination which has been amply verified under game farm conditions. In the wild, however, development sometimes takes several days longer, low^ temperatures or absence of the female from tlic nest ;i|i|i;ucmi1\ being tlie primary factors responsible for diflerences which occur. During this |)crio(l. tin' bir most of the time. On the other hand, she nuist also feed. In order to secure some idea of tlic d;iil\ routine of nesting birds during this time, two nests on the Coiuicctiiut Hill area were watched for >e\('ral days from tree blinds during June, 1932, shortly before they were due to hatch. These birds sal continu- ously except for infrequent brief periods of absence, presumably to feed. These recesses usu- ally consumed from 20 to 40 miiiules. although on one occasion, the bird remained awav 110 minutes. Over a period of three days i following which the nest was broken up I the first bird left to feed each evening between 5:30 and 7:30. In addilion. she also icfl for 'M\ minutes between 4:00 a. m. and .S:00 a. m. on the third da\. llic sei'ond bird was also observed for three days, following which the clutch hatched. The first day was stormy and the female was CHARACTERISTIC OF THE FEMALE 289 on the nest constantly after observations began at 9:30 a. m. The second day she left at 1:47 p. m. for 22 minutes, which was the only absence that day. The next day. however, she was away from 9:50 to 11:50 in the forenoon, as well as for two approximately half-hour intervals between 5:00 and 8:00 in the evening. While on the nest, both of these birds spent most of their time in utter immobility. Occasion- ally they would change their positions or adjust the eggs. Once one bird picked at objects about the nest, presumably insects, but without moving her body. When leaving the nest, both birds most often walked a step or two and then flew, often some- what noisily. In returning, the birds invariably walked, although they may have flown to within 15 to 20 yards. A bird, observed by Smyth''^ flew to a nearby tree from which she inspected the vicinity before alighting and walking to her nest. Another characteristic of incubating grouse is the formation of much enlarged droppings or "dockers". These are sometimes found in the vicinity of nests in the direction the female habitually travels in leaving. Nesting Tolerance Customarily a tolerant species, there .seems to be little conflict between nesting grouse and other nesting birds, in New York at least. The remaining gallinaceous species to be found have occupied, in general, different habitats or are so rare that they seldom nest nearby. No evidence of competition between grouse and the manv small, woodland birds has been noted. In localities where woodlot and farm environments overlap, however, the pheasant frequently lays eggs in nests of the grouse. But. despite such parasitism, the latter usually proceeds to incubate, often with the result thai the jiheasant eggs hatch first, thereby sealing the doom of her own clutch. .4^fi U. Smith GROUSE NEST CONTAINING SIX PHEASANT EGGS 290 GENERAL HABITS Of 2,016 nests examined from 1930 to I9-i2, 11 contained pheasant eggs in addition to those of the grouse. There seems to be little question, however, that such instances are the result of a parasitic tendency on the part of the pheasants involved rather than competition for the nesting site. Another report'^'' tells of nests of a grouse and a domestic hen that were but four feet apart. This account also goes on to relate that the grouse nest was broken up, following which the grouse laid several eggs in the hen's nest. Analysis of the relative proximity of different nesting grouse has revealed some wide varia- tions. The examination of a large number of nests over the last 12 years has sliuwn that female grouse seldom nest within 100 feet of each other and that intervening distances of over .500 feet are the rule. Nevertheless, scattered instances of two or more nests quite close together have been recorded. In one, three nests were found all within 50 to 100 feet of each other. In another, two nests were 50 feet apart, and in a third instance the distance was 75 feet. Furthermore, a number of grouse nests have been found in which there was every indi- cation that two hens contributed to the clutch present. It seems probable that nesting grouse exhibit little, if any, intolerance toward each other. The usual scattered distribution of nests is more likely the result of a lack of need for any greater concentration. In those cases where grouse nests have been found close together, it is possible that both females mated with the same cock. Desertion Occasionally, female grouse desert their nests. Such action is usually the result of undue interference with the bird by some outside agency. Rarely is it due to a cessation of the maternal instinct. Desertion from the former cause probably occurs most frequently before any eggs have been laid. On both the Connecticut Hill and Adirondack study areas, perfectly formed nests have been found in which eggs were never laid, though, in some cases, the female was flushed from the spot. Later, a nest was often found a short distance away iti w'hich a full complement of eggs was deposited. In one case, the second nest was directly beside the first. If the provocation is strong enough, however, desertion may occur at any time, although the lirobability decreases markedly as incubation progresses. It is probable that desertion, due to a cessation of the maternal instinct, occurs largely in connection with late rcnesting birds. One bird incubated until July 5 before giving up. In this case, however, examination of the eggs showed them all to have been infertile. It is, of course, evident that the death of the female away from tlic nest would give the impression of desertion. Thus, in analyzing such records, any error will tend to increase the percentage of frequency. In any case, however, the incidence of desertion*, after eggs had been laid, has been almost negligible among the nests examined during this Investigation. That the birds will endure a considerable amount of interference, especially during the latter part of the incubation period, without deserting, is evidenced by the experience of the authors in trapping nesting birds for marking purposes. Such trapping operations were carried on at the nest site during the last few days prior to hatching. Of 128 females so handled on the Con- necticut Hill area, only eight or 6.2 per cent deserted. In this connection, one bird, which accidentally broke its wing in a trap, nevertheless re- turned to its nest, hatched the eggs and successfully reared the resultant brood. Although crip- • See Chaplet \\\. p. S27. CHARACTERISTIC OF THE FEMALE 291 pled during most of the summer, it was finally found to have regained its ability to fly by late August. Renesting Since the latent ability of a female grouse to produce eggs* usually exceeds the number comprising her first clutch, hens, whose initial attempt is broken-up, often try again. Each year, while the great majority of the successful nests in a locality hatch within a few days of one another, a few are incubated much later and hatch on widely separated dates. Similar evidence is also found in the occurrence of broods which are much younger than the average. For this purpose, a new site is invariably chosen. And the number of eggs laid is normally less than before, averaging 7.5, although as many as 11 have been recorded. The probability of a second effort of this kind is greatest when the first clutch is destroyed before it is complete or before incubation has progressed very far. That it may. however, occur considerably later was demonstrated in 1940 when a brood of six chicks, not over three days old. was found July 24. The female of this brood must have begun to lay her final clutch about June 16. On another occasion, a renesl hal( hed July 1 or 2, and a third hen incubated until July 5 before abandoning a clutch of infertile eggs. There is no indication that third attempts are ever made. Lack of Second Broods In the region covered by this Investigation, individual female grouse have but one brood a year. The same is true throughout the range of the species, so far as is known. Normal first nests hatch in late May or early June and the subsequent broods regularly remain to- gether until September or October. The discovery, now and then, of late broods has led to the misconception by some that second broods were produced. Actually such observations merely represent birds which re- nested^ after their first clutch of eggs had been destroyed. Relation to Brood Bold and timid at once, the grouse is one of the most courageous of the "mothers of the wild", especially during the early days of the brood period. Defense of Chicks The impulse to protect her brood from harm becomes stronger and stronger during incu- bation and reaches its peak at the time the eggs hatch and immediately thereafter. Should she be molested at this time, she may either confront the intruder belligerently or endeavor to lure him away by a ruse. While the chicks or hatching eggs are still in the nest, the female most often resorts to the broken-wing "stunt" in an effort to divert attention from them. Being highly precocial, the chicks leave the nest within a few hours after hatching. For the next week or ten days, when danger threatens, they immediately take advantage of whatever shelter is at hand and "freeze". On such occasions, for the first three or four days, the female gives a warning call to her brood and at the same time faces the intruder "with blood in her eye". Ruffling out her feathers and half spreading her wings, she usually utters a loud squeal, apparently to distract his attention. Again she may rush at him. hiss- ing menacingly as she comes. Birds have been known to actually fly into the faces of per- * See Chapter VHI, p. 360. A See discussion under Reneating above. 292 GENERAL HABITS sons as well as animals. After a moment or two of this, which the nest site. I'ntil the chicks begin to assert their independence in late summer, the female shepherds her brood, drawing their attention to desirable food items and keeping a constant watch for danger. During the first two or three weeks and occasionally thereafter she hovers them at night and protects them during stormy, chilly weather, l^p until they have commenced to fly, she also broods them at intervals through the day. During the latter [)art of the summer, the faniilv group spends many hours sunning and dusting, mainly during the niiddle of the day. On such occasions, llic hen is ciiniparativch inacli\c wiiilc llic chirks |)l;i\ almul her. Kxpericnce with broods held under artificial conditions iridicalji's that the degree to which the female "talks" to the chicks as they move along is of major importance with respeit to her ability to control and guide them. An interesting observation in this connection is that, when a brood is disturbed, the female often seems content to move off with onl\ two or three chicks in spite of what may he ha|)pening to the rest. Yet. if the whole brood is threatened, she will display considerable solicitude. As the season jirogresses and the chicks develop, the role of llic Icrnalc as guide and protec- tor becomes less im|)ortanl. By Sc|)leinl)cr. the youngsters have l)ecome essenlialh adult and. about the middle of the month, the family grouj) usuall) begins to break up. CoiilKil i)j Cliirk.s During the earlv weeks before th<' chicks have become strong lliers. the female soon returns after a brood has been flushed. If danger is still |)resenl. she warns the chicks to remain hidden as well as (piicts aoN that begin to peep. The lunod mar lie kept thus concealed for several hours. The female, meanwhile, circles around or somelinics Hies to thi" low branch of a tree. contiiuialU r<-peating her warnings to the chicks. \\ hen she has satislicil herself that the coast is clear, she calls the chicks which quickly gather fmni their hiding |daces. The fdllowiii''. from the field note- of June 12. 1 ').'^2. is lllii^trati\ c : CHARACTERISTIC OF THE FEMALE 293 "Brood 4-III was flushed. The female was chased away to a distance of 200 feet and one of the party concealed himself in a thick white pine about 20 feet from the ground. Within ten minutes, a chick began peeping 30 feet from the base of the pine. Then the female was heard giving an alarm note about 40 feet on the other side of the pine from the chick. The female approached rapidly on the ground, as could be told by following her continuous soft clucking. Every moment or so between clucks she would give an alarm note. When she was right under the pine in which the observer was concealed, she flew to the limb of another pine about 10 feet from the ground. For two or three minutes she alternately clucked and warned the chicks. The one chick stopped j)eeping. When the female was sure no danger was near, she gave a series of low clucks and in two or three minutes all of the chicks collected around her. Immediately they moved swiftly off to the east, with the female clucking softly." That the female is not indispensable, at least to older broods, has been demonstrated on numerous occasions by the chicks moving off individually and collecting themselves. Ofttimes, also, older chicks will endeavor to silence other chicks which start peeping. Don hie Broods On a inimber of occasions double broods have been iMirountered. usuallv with both females present. In most cases, these a|)|)ear to rcprcsctil mcrcK chance mi'ctings. Ibi- fdllduiiig is from the field notes of June II. 1932: "Two broods were flushed logcllicr in Section 1. Ihc female (if one brood was marked, the female of the other was not, so that they were easily dislinguishable from one another. The females were chased away while two observers concealed themselves in se|)arate trees about 30 feet from the ground. In about 1.5 miimtes, a chick began to peep. Soon another was heard. The chicks of the two broods had become all mixed together, but were apjiarenllv very easily separated by the females as follows: First the marked female was heard about 100 feet away. She warned the chicks, then came up swiftlv and passed directly under one of the observers. All of the time she was giving alarm notes. After a few minutes she began to call softlv. Her own chicks came from all directions, some ]ieeping loudly, and they all moved off quickly to the north. Five minutes after the\ left, we heard the other female, who seemed more wary than the marked bird. She passed under one of the observers and flew to a low tree. Then she called her chicks together. It was about 20 minutes from the time the marked female was flushed until she had collected her brood and moved away. It took the other female about 30 minutes. There seemed to be lui iroiiMe in separating the chicks." There is some indication, however, that, later in the season, broods mav sometimes merge permanently, especially if the female of one has been killed. In other cases. or])haned broods have succeeded in maintaining themselves as a unit. Another interesting observation involves a female, whose nest was known to have been broken up just before hatching, which was later flushed several times over a period of a month with a brood of four chiiks. The identity of the hen was positive because she was marked with colored tail feathers and the chicks were of normal age. In this connection, females which have lost their entire brood, especially during the early part of the season, usually continue to act quite broody for sometime thereafter when flushed. Guidance Juveniles of any animal are. to a considerable degree, dependent on adult guidance in "learn- ing the ropes" of their environment. This is particularly true of precocial species, such as the 294 GENERAL HABITS grouse, where the chicks leave the nest within a few hours after hatching. They must be shown suitable sources of food; they must be protected during adverse weather; they must be taught how to avoid the many dangers constantly threatening them. Since the male grouse takes no part in reproduction beyond the mating stage, this task devolves upon the female. During the period prior to flying, the chicks travel in a loosely-knit group with the female, seldom straying more than ten or 15 feet from her. j\t this time she maintains a considerable degree of discipline over them by means of a variety of calls and warnings. Many times broods of this age have been observed crossing country roads with the female in the tniddle of the highway and the chicks strung out in more or less single file from one ditch to the other. Yet among broods held in large natural enclosures the chicks often seemed to lead the hen, spread out fan-wise before her. With respect to feeding, the female .seems merely to guide them through good feeding terri- tory and, by example, to indicate choice foods. She apparently does not scratch for them; neither does she feed them directlv. When danger threatens, she warns the chicks to hide and keeps them quiet until the cause for alarm has passed. As the chicks become older, they more and more assert their independence, and the degree of guidance from the female diminishes. As her control over them lessens, the group ranges farther and farther afield. Hand in hand with this, an increase occurs in the juvenile mortal- ity rate, which has largely leveled off following its initial precipitousness of the first three weeks. But, by fall, the survivors have acquired the ability to take care of themselves and the female's responsibility has been fulfilled. CHARACTERISTIC OF THE RROOD The chicks, too, even though indulging in activities common to the adults, exhibit certain distinctive characteristics. Area Traversed From the time of leaving the nest until they break up in the fall, broods range to varying degrees in quest of the necessities of life. There is nothing, however, to indicate that juvenile grouse recognize territorial limits in the sense that this term has been applied to the adults. Nevertheless, the area covered undoubtedly reflects the nature of the female's territory as well as her individuality. It is impractical to attempt to follow grouse broods from day to day. Yet it has been possi- ble to contact most broods from two to four times a month, during the summer, on the study areas. In this way. a reasonable estimate of the extent of their wanderings can be made. The identity of individual broods can usually be determined on the basis of location of contact, runnber of other broods in vicinity, number of chicks, age of chicks, habitual actions of female and artificial markings of female. Unless the female is marked, however, contacts of broods which have traveled unusual distances will often be incorrectly diagnosed. Brood "territories" estimated in this way have varied considerably in both size and extent. They have ranged from between five and ten acres to over 100 acres and from relatively com- pact imits to narrow, much elongated tracts. In 10.^1. the distance between the extreme points visited by Brood l.S-I was estimated at 3.76.5 feet. Many others have reached points over half a mile from where thev started. Nevertheless, the majority confine their activities to areas of between one-eighth and one-quarter mile in extent. CHARACTERISTIC OF THE BROOD 295 The most restricted areas of this kind have been observed on the Adirondack study area. In this section, broods commonly occupy alder beds and limit their activities to the confines of such cover units to such an extent that they rarely venture beyond their borders until late summer or early fall. For example, throughout each of several summers, two broods occupied a covert of this kind comprising, in all, only about 15 acres, each keeping to its respective end of the tract. On the other hand, many broods were found in the more usual types of cover on this area and ranged over considerably larger areas during the summer. In the more open type of range of the central and southern portions of the state, of which the Connecticut Hill and Pharsalia study areas are representative, the broods tend to follow the woods edges over much greater areas, although instances of restricted movement have also been noted here. The average, however, is about 40 acres. Examination of the courses followed by broods, as revealed by successive contacts, demon- strates the significant influence of woods edges, the borders of clearings, .slashings, woods roads and the like on the territories chosen. In areas of disconnected coverts, a large propor- tion of these areas are located about the periphery of the existing woodlots, many extending for considerable distances along the woods edge, but pciiclratiiig the interior hardly at all. In tracts of continuous forest, however, where openings arc c(ini])aralivcly small, the broods, in following their borders, usually remain within more conipact limits. This is also true of broods which choose the vicinity of small openings within the woodlots of disconnected coverts. Observations indicate that food is a more important factor than shelter in governing brood movements* although even young broods are often found in conifer patches. Usually they make their way at once to situations where there is an abundance of low, shrubby vegetation with its accompanying po|)ulatioii of insect life. In such situations, the sunlight, in addition to being a necessary element of the conditions involved, aids in keeping the chicks warm and in drying them ofF after a storm. As the season advances, suitable food continues to become available in much the same types of cover. Then, in late July, the raspberries and blackber- ries connnence to rijjcn and a noticeable movement to the brier j)atches takes place. There seems to be no tendency for broods to olijcct I(j the presence of another brood on the same area. Many cases of over-lapping routes luuc been recorded. In fact, as noted else- where.^ two broods have been found to actually merge and subsequently separate again. That, at times, even quite young broods will cross appreciable open areas has been demon- strated on several occasions. For example, one brood less than two weeks old, the female of which was marked so that identification was positive, was found to have crossed 600 feet of open field to a separate patch of woods from that in which it nested. This brood later returned to the original covert. Several other similar instances have been noted where broods which hatched in small separated woodlots have traveled to a larger tract and. often, back again, crossing open fields of several hundred feet en route. This, however, is not a frequent occur- rence. Hiding The ability of grouse chicks to "vanish before one"s eyes", so to speak, has long been commented upon. Contrary, however, to popular belief, they do not grasp a leaf in their feet and roll over. Their technique is merely to squat as low as possible and "freeze". In this » See olso Chal.tors HI nnd IV. A See disriission iindcr Relation of Female to Brood, p. 29:i. 296 GENERAL HABITS way. they may often be in plain sight and yet be ahnost invisible, due to the blending of their plumage with the sunlight and shadow of the forest floor. It is true, however, that they fre- quently scuttle under a leaf or other protecting object before "freezing". Although this practice is chiefly characteristic of chicks before or for a short time after they ha\e learned to fly. it is often utilized later in the season. Furthermore, the same method is employed by older chicks at the point where they settle down after being flushed. While in hiding, the chicks usually remain silent for upwards of ten to 15 minutes but then frequently become impatient and commence to peep so that the female, which customarih remains in the vicinity, must quiet them with warning calls*. Among other chicks, the effect of this sometimes wears off and continued peeping occurs. At such times, chicks have been seen to leave their hiding places and start out bv themselves. Usuallv. however, the danger which caused them to go into hiding passes before such a state of unrest is reached. Older chicks, on the other hand, have been known to steal away, silently and unnoticed, under the very nose of an observer watching for them. Roosting During the first few weeks, the chicks are usually hovered by the female at night. Experi- ence with broods held in large outdoor pens offering a variety of cover indicated that it may be common practice for hens to hover their chicks in the open rather than within the woods. Rroods but slightlv over two weeks old. however, have been found roosting in the low branches of trees and shrubs. Later, this habit becomes customary, although well advanced broods frequently roost on the ground, usually in the shelter of a conifer. Young broods roost in very closely bunched groups but as the chicks become older and more independent their association becomes progressively looser. Feeding Habits Thrfiughoul the brood period, grouse chirks spend most of their waking hours on the ground. Feeding is their princijial activity although, for a lime after hatching, brooding is frequent and during the latter part of the summer much time is spent dusting. • Sec ChnpliT M. p. >W. CHARACTERISTIC OF THE BROOD 297 During the first two weeks, insects constitute a large proportion of their food, while later vegetable material is predominant. Young chicks, especially, are a hive of activity, constantly picking amongst the ground litter or chasing insects through the low-growing vegetation. In the course of such pursuits, they often flutter off the ground a bit. Older chicks frequently jump eight or ten inches in the air. aided by a wing flap or two, in order to get some particular morsel out of normal reach. They scratch very little with their feet and the hen seldom passes on to a chick a food particle she herself has picked up. She may, however, sometimes call the youngsters to her and encourage them to eat some special delicacy she has found. As they proceed, the chicks sample everything, both animate and inanimate. Bits of wood and other debris are apparently taken just as readily as a juicy insect, although they are usu- ally dropped again. Movement, however, definitely attracts their attention. This is probably one of the chief reasons why ants are eaten in such large numbers. X^lien a particularly in- teresting new object is encountered, the chick will stretch its neck, inspect it with one eye and then, turning its head, give it the once over with the other eye. Activities of this kind are, naturally, very difficult to observe among wild birds. At the Catskill Experimental Station, however, several broods were hatched and reared by grouse mothers on the ground in large open enclosures affording an abundance of typical cover as well as excellent opportunities for observation. These broods usually began their day's adven- tures shortly after sun-up. Even rainy weather did not long delay the start. For the first week or ten days, before they began to fly, the chicks were brooded at night by the hen. In the morning, she would get up. the chicks tumbling down from under her wings or emerging from some other retreat. Immediately, she would move off some ten feet and wait while her brood, spreading out fan-wise as they chased bugs and beetles or drank dew from blades of grass, worked up to her. On warm days. u|)wards of half an hour might be spent in this way, but, when the day was cold or rain\. the chicks often returned to be brooded again after only five minutes. Thus, throughout the day, periods of feeding alternated with periods of brood- ing. Seldom, however, would all chicks be under the hen at one time, individual birds being continually on the move. As the birds grew older and commenced to fly. they usually roosted at night and brooding became discontinued altogether. For the remainder of the summer, while feeding might be indulged in at any time, it was less common during the middle of the day which was devoted more to sunning and dusting. Dust baths, however, are often associ- ated with ant-hills and there is every indication that large numbers of ants are picked up on such occasions. Dust Bathing What has been said under this topic with respect to adults applies equally here as to func- tion, sites chosen and materials utilized. Chicks have been seen dusting when only about two weeks old but the general onset of this activity is usually a week or so later. From then on, it is common to find dust baths comprising one large depression flanked or surrounded by several smaller ones, where a female and her chicks have had a family dusting party. Gregariousness In plotting the travels of individual broods it is not unusual to find that the areas covered by two or, occasionally, three such groups have overlapped. On the other hand, it is unusual 298 GENERAL HABITS to find both broods at the same point at the same time, although this situation has been en- countered on several occasions. In these instances, both females have usually been present and have subsequently gone their respective ways with their own broods. But in one case, two such broods were again found together the following day, while two others were found together a second time after an interval of two months. It seems probable that such occur- rences are largely accidental. Now and then, however, particularly late in the season, a group of orphaned chicks has, apparently, joined another brood permanently. '''''^^X^^^^: CHAPTER VI INFLUENCE OF WEATHER by Walter F. Crissey WEATHER AS A DIRECT DECIMATING AGENT WEATHER AS AN INDIRECT INFLUENCE During the Nest Period — During the Brood Period — During the Adult Period COVER TYPE USE AS AFFECTED BY WEATHER WEATHER AND FLUCTUATIONS IN GROUSE POPULATIONS ^ Summary Weather is a basic influence on grouse and other wildlife species through its effect on the environment in which they live. (p. 300). Its effect is largely indirect and specific relationships are therefore difficult to determine. (p. 300). Direct losses from weather conditions seem negligible in New York, but on one occasion a severe cloudburst and flood before the chicks were six weeks old apparently took a heavy toU. (p. 301). Although the onset of the reproductive season is primarily controlled by the progressively increasing length of daylight in the spring, minor variations in nesting dates from year to year seem related to the average minimum temperature during early April, (p. 303). There is apparent correlation between temperature and precipitation during the three weeks immediately following hatching and the brood mortalitv occurring during the latter part of the summer, (p. 303). The degree of over-winter loss among adult grouse seems to be affected to some extent by the severity of the weather during March, (p. 304). There appears to be some degree of agreement between the occurrence of low temperatures during March and June and periods of grouse scarcity which have been recorded in the past. (p. 306). 300 INFLUENCE OF ff FATHER Weather, in its various forms, is one of the basic influences affecting the character of organ- ized life throughout the earth. During the planets formative stages. i)recii>itation and frost action motivated the forces of erosion which gradually broke down the original rock of the earth's crust enabling primitive plant life to gain a foothold. Rainfall, as well as melting snow and ice, also gave rise to the oceans in which, under warm temperatures, the earliest forms of life developed. Throughout succeeding eons of geologic time, these same forces have been instrumental in establishing the vast array of rock formations and soil types which are funda- mental in supporting the vegetation upon which animal life so greatly depends. Then, too, conditions created by weather have been directly responsible for many of the major changes in life over the earth's surface. Today the distribution of plants and animals, as well as the nature of many of their characteristics, rests to a large extent upon their reac- tions to conditions controlled by weather. Animals inhabiting these regions, therefore, have become physiologically adapted to certain conditions. When their range reaches an area where the environment is excessively severe, they cease to occur. Similarly, within its normal range, the abundance of a species from year to year may be affected by variations in weather. Despite its universal significance, however, knowledge of this factor is still quite limited. The science of meteorology has advanced to the stage where short-time weather forecasts can be made with fair accuracy. But prediction of the effect of such conditions upon the growth and abundance of plants and animals is largely speculative. A major reason why existing knowledge with respect to wildlife is meager lies in the com- plexity of the manner in which weather usually exerts its influence. In the first place it com- prises a number of components: temperature, precipitation, sunshine and humidity, to mention a few. But more important are the high variability of these forces, the rapidity with which they fluctuate and the fact that their pattern is constantly changing. Furthermore, the indi- rect and often delayed nature of their effect on an animal tends to obscure the actual relation- ship. Another reason involves the applicability of existing weather data to wildlife problems. The standard observations of the Federal Weather Bureau afford an excellent index to the general trend of the various climatic elements. But such records, even those of the nearest individual station, usually differ from the specific conditions experienced on local areas where wildlife studies are being conducted. Therefore, while it is possible, in many cases, to recognize cor- relations between weather trends and the behavior of animal ])opulations, it is usually much more diflicult to determine the mechanism responsible*. Weather, with respect to grouse, has long been considered an important factor affecting its abundance from year to year. Many observers have suggested cold, wet conditions during May or early June as a primary cause of scarcity the following fall through in(rcasc kind will undoubtedly use up less energy in combating heat loss than luu- with- out such protection. This mav explain why grouse are apparentlv capable of remaining in roosts of this sort for considerable |)eriods without food. To Mini up this discussion of its indirect influence, weather may be considered primarily as a factor conditioning the susceptibility of grouse to more direct decimating agents through sapping their vitality. Actually, death is usually due to |)rc(Iaticin or disease. * Statistical nnAJvait ha« tliuwD ihem to be highly ■ignihcanl. A Sn Chapter U, p. 91. t Sec Chapter \ . p. 270. COVER TYPE USE AS AFFECTED BY WEATHER 305 COVER TYPE USE AS AFFECTED BY WEATHER In grouse management, certain cover types, under various circumstances, are important. If particular types are necessary for survival under adverse weather conditions, then they should form a part of the management plan. This problem was approached by attempting to determine whether use of any type of cover was more particularly associated with one form of weather than another. Analysis of the locations in which grouse were flushed under different conditions, however, for the most part, proved inconclusive. Nevertheless, both adults and broods seemed to frequent the heavier types of cover, conifers in particular, to a greater extent during rainy weather, while the more open types, such as second growth hardwoods and overgrown fields, were used more during dry weather. In general, differences accompanying the change of seasons influence the use made of many types of cover to a considerable extent. But the issue is confused by the fact that the character of the cover t) pes themselves changes with the seasons due to the growth and loss of leaves from deciduous trees as well as the production of nuts and other fruits. Probably the most definite relationship with respect to weather occurs during the winter. In the Northeast, areas which lack sufficient coniferous shelter seldom maintain a good grouse population. Other types are necessary to grouse but not particularly from the standpoint of weather*. WEATHER AND FLUCTUATIONS IN GROUSE POPULATIONS The fact that weather conditions may have an influence on brood and adult survival among the ruffed grouse has been shown in the foregoing portion of this chapter. It has been noted 0 — c\i r) ^ irt * ffi a^ o> o> oi c> ^ 01 O S ID CO (O y the environment. Thus the shaded thermometer at a "weather station" may regis- ter 20 degrees below zero, but if a grouse occupies a protected nook on a sunny south slope during the dav and resorts to a snow-roost at night, it has succeeded, to a considerable extent, in minimizing the effect of that temperature. The problem is complex, to say the least. * The valiip used par\\ yrjir ronftistpd uf llir mint of the ili-viiiiiuii^ fnini the »tnndatil mruri of ihr uveragc lemperaturrt for Marrh anit June. CHAPTER VII PREDATION By Robert W. Darrow COMPETING INTERESTS ROLE OF PREDATION Effect During Each Life Period — Nest Period — Brood Period — Adult Period — Predator Fluctuations — Buffer Fluctuations — Predator Pressure ■ — Grouse- Predator Ratios — Vulnerability — Significance of Predation APPRAISAL OF THE VARIOUS SPECIES Grouse Predators of the Northeast — Rei^tionships During Each Life Period • — Nest Period — Brood Period — Adiill Period — Net Effectiveness Throii.hoi t the Year — Food Habits in New York — Fox — Weasel — Skunk — Raccoon — Mink — House Cat — Great Horned Owl — Barred Owl — Snowy Owl - Goshawk — Cooper's Hawk — Sharp-shinned Hawk — Red-tailed Hawk — Red-shouldered Hawk — Marsh Hawk — Interpredator Relationships THE PROBLEM OF CONTROL Experiments Conducted — Effect on Predators — Results During Each Life Period — Nest Period — Brood Period — Adult Period — Effect on Productivity &<. SUMMARY Predation is a fundamental relationship of living organisms funt iioiiing. through the removal of surpluses, to keep their numbers within limits compatible with the habitat they occupy, (p. 308). Although increases in a grouse population cannot be accumulated year after year, predators or no |)redators. control of foxes and weasels may be useful in producing a greater fall surplus on areas where such a crop will be utilized for sport, (p. 331. 350). \m a\crage of approximately 39 per cent of the grouse nests observed by the Investigation have been broken up. mainly by predators, (p. 311). Foxes, the principal grouse nest predator, have broken up more nests when buffers were scarce than when they were abundant, (p. 3151. Only about half the usual brood mortality seems attributable to predation. (p. 317). 308 P RED AT I ON Adult losses have averaged slightly over half the fall population each year, some 80 per cent of which, on unhunted areas, succumb to natural enemies, (p. 319). The numher of adults lost has varied directly with the fall population level, fp. 320). Great horned owls, the major predators of adult grouse over most of New York, have in general taken more grouse as buffers became scarcer, (p. 322). Resident predators have not fluctuated greatly in abundance on the study areas. I p. 323). Buflfers have exhibited marked fluctuations, some approaching a cyclic pattern, (p. 32,S). The degree of activity among foxes, and thus their pressure on grouse, has varied inversely to the abundance of buffers, (p. 328). Grouse predators include a wide variety of species ranging from tiny shrews to the wulf and great horned owl. (p. 332). Foxes and weasels are the chief predators during the nest period, (p. 334) : Cooper's and sharp-shinned hawks among the chicks, (p. 336) ; and the great horned owl and fox among the adults, (p. 337). In no instance do ruffed grouse constitute the staple food of any predator. I p. 338^. Predator control experiments have been effective in reducing nest losses, but have had less effect on adult mortality and little or none w ith respect to the chicks. I p. 319 ) . Predator control cannot be depended upon to result in a continued increase among breeding populations of grouse, (p. 350). "The large fish eat the small fish; The small fish eat the water insects: The water insects eat plants and mud." So it is with terrestrial and avian species as well. The fox feeds largel) on rabbits and mice which in turn live on vegetable material. The Cooper's hawk preys on small birds which eat insects which subsist on i)lants or decaying animal matter. Predation is one of the fundamen- tal relationships of living organisms. Hut because instances are easily obser\f(J and lieu|iul,ilioii surpluses. That it represents » S (,K()ISE NEST SHO\MN(, FEATHKKS l)F FEMALE WHICH WAS PROBABLY KILLED BY A GREAT HORNED OWL AND SEVERAL EGG SHELLS GIVING EVIDENCE OF HAVING BEEN EATEN BY A RACCOON But, while the degree of loss over a region as a whole may average very nearly the same for successive years, local areas often vary more widely. The record for Connecticut Hill is an example (figure 24). During most seasons the number of such local areas above average tends to balance those below. Now and then, however, as in 1940, a majority lean the same way. 314 PREDATION In either case, hfiwever. predation lias heen refiiilarlv responsible for nearly 00 per cent of the nests which failed*. Beyond this, predators. |)articularly the fox", often filch eggs from clutches, the remainder of which hatch successfully. In other words, over 35 per cent of the j)otential grouse crop*^ each year over the State I outside extensive forest areas) has thus been destroyed while still in the egg stage. \^'ithin the Adirondacks the latter figure has been somewhat lower since total nest losses have been less. The same is probably true of extensive tracts of timber in the Catskills although data are lacking. Predation of grouse nests niav take place at any time from the laving of the first egg to the date of hatching. In general, however, the rate of loss has been greatest toward the end of the incubation period. While experiments with bird dogs have indicated that sitting grouse emit little scent it has been observed that, in many cases, as incubation progresses the eggs become more or less fouled. Under such conditions sufficient scent to attract predators may be given off, especially when the female is away from the nest. Furthermore, there is some evidence that, where a high breeding population exists, certain animals, particularly the fox. often happen on enough nests as the season progresses to associate the flushing of a bird with the possibility of dining on "omelet". z < 2 O u. z o \- > UJ I- o a. ui loo I a. SO So L Buffers (robbi4a , rr»icc and sWrews) 200 l-J l~f 1933- S* 1934-36 I935-34 193^-37 YEAR 1937-38 1936-39 I939-40 1940-4-1 ZOO KIGURK 2.5. INTERRELATION OK GROUSE NEST M(IKTM,n\ \\ 11 H FOX ACTIVITY AM) Bl FFER ABUNDANCE — CONNECTICUT Illl.I. ARK\ I').'U-I*>H l'.\cii though in most \ears during the period sliidicd the average nest mortality for the Stale has not differed greatly, the fact that it varies more widely on local areas is of interest to the si)ortsinan and game manager. Since ])rcdatioii is the primary cause of such losses, predator pressure must vary similarly. That it does vary has been discussed elsewhere^. * Scr ('.ha|>Ii'T \U. [1. SJ7. (or iliiiriiiittinn ii( othrr rauiir» in\iil%'cd. A Either ihe red (oi (Vutpes lulva) or the gray fox (Vrocyan cinertoarirnttui) t Apptirt only to nrnln arliinlly rnlabliiihril. X No cyclir drrlinr hnn been rxprrirni-r<] Hiirinc ihr ln\r«ligalion. § See diKcuatiun o( Predator Prc»«urc, p. 3^7. ROLE OF PRE DAT ION 315 together with data indicating an inverse relationship between the abundance of small mam- mals and the degree of activity among predators. In figure 25 is shown the correlation between the degree of activity of foxes from year to year and nest losses during the corresponding season on the Connecticut Hill area. The simi- larity of the two trends is striking. Apparently they are directly related, although in 1941 some other influence became controlling. Thus under the conditions studied foxes must be considered to have been a major factor in determining grouse nest failure, and, in turn, the pressure exerted by them has been greater when small mammals have been scarce than when they have been numerous. Reference to figure 24 shows a high nest loss on Connecticut Hill in 193,S. While com- parable data for buffers are not available prior to 1934 general records indicate a lower buffer abundance during that year than at any other time except the spring of 1937. It is not surprising that a substantial proportion of the grouse nests attempted each year should l>e found by predators and destroyed. A ground neslcr the female commences to lay during the latter part of April and does not bring off her brood until about June 1. Thus for a period of some six weeks or more the eggs are in constant danger of discovery by the numerous animals which would welcome such a ( hange in diet. At this season these species spend considerable time hunting, either to recuperate from winter privations or to supply their growing young of the year. True, the coloration of the female when on her nest blends remarkably with that of the forest floor. But when she is not sitting the eggs are usually more conspicuous, since it is only accidentally that leaves tend to camouflage them*. The Brood Period Commencing with the hatching of the eggs, the brood period has been considered to extend through August 31. Unlike nests, however, grouse chicks do not "stay ])ut". Sur\ ival records were therefore secured largely from those areas where summer census surveys were regu- larly conducted, namely, the Connecticut Hill and Adirondack areas. An average of 42 and 12 broods were studied each season on those tracts, respectiyely. Table 37 presents the per- centage mortality"^ observed each year. TABLE 37. BROOD MORTALITY RLC.ORDLD ON CONNECTICUT HILL AND ADIRONDACK STUDY AREAS— 1930-1912 Area Year 1930 1931 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 1942 Aver. Coiiuectirul Hill Adirondack 57.5 70.0 5t.8 58.3 76.7 78.4 51.4 50.8 80.8 88.3 54.2 41.8 55.9 60.0 62.4 72.0 63.0 57.0 57.9 57.0 59.5 64.0 77.6 55.5 63.2 60.9 It is evident that losses during this period have been proportionately greater than with respect to nests. There has also been a remarkable similarity both in trend and degree from year to year on the two widely separated areas. Yet an adequate appraisal of the relative importance of the various factors' involved represents one of the most perplexing problems the Investigation has encountered. * See Chapter V. p. 288. A .'^ItlHMigli teriin'ii hrmxl riiuitalitv. percentages refer to the aggregate loss of chicks, t See Chapter Xll, p. 528. 316 PRE DAT I ON Dead chicks have proven extremely difficuU to find. In spite of heavy losses during the first few weeks after liatching the most intensive field work has revealed the remains of but very few. Not until nearly the first of August does one begin to encounter tell-tale piles of feathers. This means little, however, for while it is true that inanv predators devour young chicks whole, it is equally true that the carcasses of indi\iduals of this age which have suc- cumbed to other causes are quickly obliterated by maggots, carrion beetles and similar agencies. In fact, after only two days, three specimens of this kind which had been placed on the ground in the woods for observation could only be identified by the stakes to which they had been fastened. Direct determination of the degree of predation at this season is thus impossible, and it has been necessary to approach the problem indirectly. Throughout the Investigation the total brood mortality has been measured each summer on the two major study areas. On both, its course has been remarkably similar, averaging slightly over 60 per cent by the end of August. Analysis of the progressive rate of decline indicates that the curve drops somewhat more steeply during June than it does thereafter. In fact by July 1 in an average year roughly half of the total brood mortality has already occurred. Connec+icu+ Hill I930-4-2 >- I- lo zo Adirondack 1332,-41 Research Cenfer (wildeqga) 1939— 4-Z 30 O I- Ao Z o a. a. 50 60 TO 2o 30 M> 50 GO lO June I ie 15 Ju lu I July 15 DATE AuQ. I \ug 15 AuQ.3l FIGl'RK 26. AVERACE PROGRESSION OF CKOISE BROOD MORTAIJTY ON CONNECTICLT HILL AND ADIRONDACK AREAS AS COMPARED WITH THAT OF CHICKS H.\TCHED FROM WILD EGCS AT THE RESEARCH CENTER There seems little doubt that many of these chicks are consumed by predators. Yet (■x])erien(c with grouse at the Research Center strongly indicates that predation is not the basic cause of these early season losses even though in many cases it may be the final execu- tioner. A striking parallel has been observed among chicks hatched artificially from wild eggs (figure 26), although here the losses have been somewhat greater. Among the latter, while it likewise remains largclv urisii!\cd. the cause is cci lainU not predation. This similarly suggests that the chief causes of these early losses are pci iiliar to this age and that predation is secondarv. This conclusion is further su|)ported by the fact that experi- mental predator control failed to alter the pattern. ROLE OF PREDATION 317 On the other hand, losses experienced during the rest of the attributable to predator activity. On the game farm mortality old is negligible but in the wild moderate losses continue. The more and more independent of the female with the result that vulnerable to attack. The principal predators at this age, the hawks (Accipiter cooperi and A. velox). were least affected by th even here, there is some evidence that variations in the degree of may often be traced to weather conditions*. summer seem more directly after the birds are six weeks chicks at this time become they probably become more Cooper's and sharp-shinned e control experiments. Yet, loss during July and August FEATHER REMAINS OF GROUSE CHICK KILLED BY A HAWK PROBABLY A SHARP-SHIN Considering all the data at hand it appears that upwards of one-third of the chicks hatched, or a little over half of the total lost during the brood period in a normal season, have suc- cumbed as a result of predation. The proportion, of course, varies with circumstances. Furthermore, it represents a greater number of individuals during those years when the hatch was higher. In other words, the more birds present the more frequently they are encountered and caught. Major differences in total losses, however, have apparently been the result of other influences despite the fact that predators may ultimately devour the birds. Thus while this factor is important in determining brood survival it does not seem to have been paramount, especially with respect to the occasional years of markedly high mortality. » Sec CliniiKT VI. p. 303. 318 PRE DAT I ON Adult Period Considering birds of the year as adult by September 1. the fall season ushers in the last of the three life periods. Most broods soon break up, the fall shuffle takes place and then winter sets in. Like their parents, the young birds must now fend for themselves and "the devil take the hindmost". Between this time and the following April the final culling of the breeding stock takes place. Again data have had to come from the areas on which seasonal censuses were being con- ducted. In measuring adult mortality two major milestones have been used each year, namely, the breeding season (April), and the following September when the next year's increment comes of age. On Connecticut Hill losses during the former |)eriiid have averaged 41.6 per cent, while for the full twelve months the mortality has been 50.2 per cent. On the Adirondack area the figures have been 52.6 per cent and 57.9 per cent respectively. The low proportion taking place during the summer has been consistent on both areas. In table 38 is shown the total mortality each year as well as the size of the initial fall population. TABLE 38. ADULT MORTALITY RECORDED ON CONNECTICUT HILL AND ADIRONDACK STUDY AREAS— 1930- 19H* Area Adult grouse data Year 1930 1931 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 Aver. Connecticut Hill Adiruudack Initial September i>upulation . IVr Cent mortality Initial September iKipuIatiou. 161 14.3 276 28.9 60A 46.7 465 57.6 70 62.9 274 40.5 51 43.lt 420 52.6 73 78.1 300 ,52,0 23 30.4 311 63.0 64 71,9 273 42.8t 59 57.6 334 51.2 42 28.6 394 58.8 70 65.7 276 54.7 63 60.3 300 57.7 53 56.6 315 50.2 6T S7.9 ♦Year uidicated is that of the boKiiining of each September 1 to August 31 period, .^Katimute made in January 1M32 und therefore somewhat below September level, till l'*33 oil the Adirondaek area and in 1937 on Connecticut ilill. mortalities of 52,9 per cent and 45.8 per cent respectively were recorded by April hut during the ensuing summers more birds moved into the areas than wen> lost. The remains of a large proportion of the adult grouse lost each year have been found and brought in to the laboratory for examination arcom])anied by a description of the situation in which each was found. They have then been studied for evidence as to what the cause of death may have been. Only about half of the specimens, however, have been sufficiently fresh and complete to diagnose*. In fact in Uj.S per cent of those looked over it has been impossible to form any conclusion at all. Of tlie balance 9L1 per cent have shown signs of predation. a proportion which undoubtedly would apply to the others as well. But whether or not this was the initial decimating agent can seldom be determined directly. One must, therefore, weigh the available evidence with respect to the other important pos- sible causes of this mortality. Besides predation only two, accident and disease, warrant consideration. The often suggested relation to birds crippled during the hunting season does not enter here since shooting has been prohibited on these areas. Regarding the former a few birds have been found which had died from injuries sus- tained ill flying into some obstruction. Most of these have occurred during the fall "crazy flight" season. Yet the majority of the loss does not take place until after mid-winter. It is • See Chapter XII, p, 532. ROLE OF PREDATION 319 significant, too, that less than half a dozen instances of this kind have been witnessed in upwards of 50,000 grouse flushes observed by the Investigation's field crews. Unless dis- turbed, these birds seldom resort to reckless bursting flight. The possibility that any large number year after year bring about their own demise by crashing into trees or other obstruc- tions in their environment seems quite remote. Turning to disease pathological examinations have shown a very low incidence of conditions which might prove fatal to grouse, both on the study areas and elsewhere over the State*. Of 143 specimens collected on Connecticut Hill less than 5 per cent could have been expected to succumb from this cause. Throughout the survey on this area but 20 birds have been found dead, showing symptoms of disease. Unquestionably many other instances occurred but there is little likelihood that this factor has been responsible for more than a very minor part .of the overwinter losses recorded during this study. Thus, by the process of elimination, one must conclude thai the great majority of the specimens found each year are attributable to predation as the original decimating agent, although it is implemented in varying degrees by a number of influences affecting the vul- nerability of the birds. Unquestionably accidents, disease and other circumstances have taken a small toll — perhaps upwards of 15 per cent. Assuming that these data are repre- sentative of the total losses occurring it seems probable that predation may be considered to have been responsible for something over 80 per cent of the annual mortality among adult grouse, or in the neighborhood of 40 per cent of the fall population on the areas involved.'^ FEATHER REMAINS INDICATING THAT AN ADULT GROUSE WAS KILLED BY A HAWK OR OWL Losses during this period have differed considerably from year to year. Especially note- * See Chapter X, p. 410. A These areas were not shot over in the fall. 320 PRE DAT I ON worthy were the very low mortalities in 1930-31 and 1931-32 when grouse had not yet recov- ered from the scarcity of 1927 and 1928. In general, however, the number of birds killed during the winter has varied directly with the population density of the preceding fall (figure 271. The proportion lost, however, has been less closely correlated*. Fall PopulaVioo ~~~ Loss +o Following Fall 500n 400 CONNECTICUT HILL in Q 300 m a. zoo u to 2 z lOO n ^°° Aoo 300 zoo loo Q 5 ■'^ ^ bo a. u to 25 D I ADIRONDACK I T5 bo Z5 1930-31 31-32 3233 3334 3-4-35 3S3C 3037 37 36 3a33 39-AO 4&4 1 41-42 YEAR FIGURE 27. RELATIONSHIP OK NL'MBER OK AI>1 l.T (.ROl'SK LOST TO PRECKDINI. KALI. I'OI'rLATION LEVEL ON CONNECTICUT HILL AND M>1H()M>\. S;I2. ROLE OF PREDATION 321 Apparently the relationship between predators and adult grouse is largely an incidental one. No species depends on this game bird for any large proportion of its food. But the greater the number of birds in a covert the more frequently they are encountered and killed. It is significant that the basic effects of predation should have been so similar on the two study areas in spite of substantial differences in the composition of the predator population. For example, the great horned owl was not observed on the Adirondack area while the goshawk was a regular visitant. Specific data from the latter area are insufficient to draw conclusions as to the comparative relationships of the various species. There is some indi- cation, however, that the fox preys to a greater extent on adult grouse on this area than on Connecticut Hill. In any case the principle of compensatory predation as discussed by Errington"' seems to be corroborated. At the same time the comparative level of the grouse population is not the only factor affecting the degree of adult mortality. In figure 28 the data have been statistically adjusted by analysis of covariance to eliminate the effect of the relative abundance of the birds. lOOa ■ lOO > O iij cc loo Q lij 2^00 a. 300 4.00 40 30 j 20 I 10 o Carouse R.emciins in Owl Pellets 4-0 30 2,0 10 O 1930-31 3\-32 3233 3334- 3435 SbiC 3t37 37-3& 36-33 3940 40-4I 41-42 YEAR FIGURE 28. RELATIONSHIP OF INCIDENCE OF GROUSE REMAINS IN GREAT HORNED OWL PELLETS TO GROUSE ADULT MORTALITY AND TO BUFFER ABUNDANCE — CONNECTICUT HILL AREA — 1930-31 TO 1941-42 Other influences, such as variations in weather and predator pressure have been chiefly responsible for the differences remaining. But just what they are and how they exert them- selves is still largely obscure. Attention may be called, however, to certain interesting cor- relations. 322 PREDATION It seems sigiiificanl that these forces reached their highest level on Connecticut Hill in 1936-37, when buffer species were at their lowest ebb. Yet they were most severe in 1934-35 on the Adirondack area where the drop in abundance of hares ( Lepiis americanus) came earlier than that i!i cottontails I Sylvilagus jloridanus) on the other tract. On the Connecticut Hill area the great horned owl (Bubo viry,inianus) has been a major predator during the adult period. In figure 28 is also graphed the incidence of grouse in the pellets of this owl collected each year. There is a strong siniilarit\ between this and the deviation from average of the adjusted degree of overwinter loss. Shown also is the relative trend in buffer abundance from 1933-31 through 1910-11. It seems significant that in 1936-37, when the greatest increase in overwinter mortality occur- red, the frequency of grouse in these pellets should have been the highest of the series and should have been accompanied by the greatest scarcity of buffers recorded. Furthermore, records of the survey give some indication that 1932-33 also witnessed a low level among rabbits and mice. In 1937-38 while rabbits were still scarce but mice and shrews had increased the incidence dropped but so also did the fall population le\el and the general overwinter loss. That the pellet representation in 1934-35 was not lower mav have resulted from the fact that the grouse population in the fall of 1934 was next to the highest recorded during the study. The data suggest that the abundance of buffers, through its effect on the amount of time jjredators spend in hunting, exerts an effect on h< O Z ^^ u a. loo ZOO n 300 Xoo loo too zoo 1933-34 34ra& 3ir3corded during the eightccnlh and early iiiTiclcentb centuries grcatlv surpassed anv which have occurred since. During the Investigation the fall of 193.'i witnessed such an irru[)tion over much of New York and New England. While many eastward ino\einents were noted most of the migration was to the west. Considerable bands of squirrels were observed swimming the Hudson River and Lake Chainplain as well as a number of lakes and reservoirs. Large numbers jicrished in the attempt. On Connecticut Hill the s])ecies increased over 300 per cent from the spring of 1934 to that n has appeared relatively stable. Undoubtedly their numbers have fluctuated but there is nothing to indicate that the abrupt changes noted between successive years might be attributed to variations in breeding success or in survival. Neither does it seem likely that the increases observed have resulted from the influx of animals from surrounding territory * This in a wnocltand fippcini in its nnlivc rnng<', A Data correUied by aoalyiii u( covariance. ROLE OF PREDATION 329 since on such occasions both grouse and buffer populations have been below average. It is, therefore, more probable that the increases in tracks recorded in 1936-37 and 1939-41 were in the main the result of greater activity associated with times of scarcity among small mammals. Figure 30 illustrates this relationship. Rabbits seem to exert a somewhat greater influence than mice and shrews. Their drop from 1936 to 1937, while the latter remained constant, was accompanied by an increase in the oc- currence of fox tracks. Conversely, from 1938 to 1939, despite a sharp decrease in mice and shrews, a greater rise among rabbits was associated with a moderate reduction in fox activity. That mice may nevertheless be important was indicated in 1937-38 when their high abundance apparently offset a severe low in rabbits. The fact that fewer fox tracks were not recorded in 1934-35 when both small mammal groups as well as grouse were very abundant seems somewhat contradictory. Yet it may be that a fox will travel about so much at this season regardless. Another reason for believing that the fox track data represent activity is the fact that the number of droppings picked up by field crews over the same period has varied inversely. Undoubtedly the great horned owl and other predators react similarly although the data have not been sufficient for correlation. With respect to pressure on grouse it follows that the more time a predator spends on the move the more often it will encounter grouse and have opportunities to prey on them. The situation appears to parallel the old adage, "you won't catch manv fish if vou don't keep your line wet". Grouse : Predator Ratios An important part of any study of the role of predation is a knowledge of the complexion and size of the predator population involved in comparison with that of the game. During the Investigation, however, it has not been feasible to undertake actual censuses of these species. Nevertheless, various data have been recorded which, supplemented bv observations during the regular grouse survey work each year and by the cumulative experience of the authors, make possible a reasonably accurate appraisal of the situation on the areas most intensively studied. On Connecticut Hill, which is essentially representative of the better grouse coverts of the State south of the Mohawk River, the principal grouse predators were the fox*, weasel, skunk, raccoon, great horned owl, Cooper's hawk and sharp-shinned hawk. With respect to this area predator control experiments carried on during 1931-32 and 1933-34 afford a basis for a rough estimate of abundance for all except the hawks. The numbers taken per square mile averaged fox — 3, weasel — 5, skunk — 6, raccoon — 2 and great horned owl — 9. These data are conservative because no species was eliminated. Furthermore, they pri- marily represent the fall to spring period. During the season when young are present num- bers would usually be higher except in the case of the great horned owl. Here the great majority of the birds trapped apparently were winter visitants. Breeding populations of this owl probably seldom exceeded two pairs per square mile. As pointed out under a pre- vious topic'^ the numbers of these species have not varied greatly on this area during the years covered. While occasional Cooper's hawks often remain through the winter the sharp-shinned hawk * The ratio of red foxes to gray foxes on this area was about 3 to 1. A See discussion of Predator Fluctuations, p. 323. 330 PREDATION almost invariably is absent at this season. In migration these birds pass through in moder- ate numbers and a few remain to breed, at which time from one to two pairs per square mile have been observed. Usually the latter species has been more plentiful. Among the predators of lesser importance to grouse on this area limited numbers of mink (Mustela vison). opossum and barred owls fStrix van'a) are present the year around. Now and then wandering house cats are encountered. During the winter goshawks a|)pear irreg- ularly and snowy owls more rarelv. The Buteo hawks (red-tailed Buteo horealis. red-shoul- dered B. Uneatiis. rough-legged B. lapopus s. johanni.s. and broad-winged B. plalvpterus) and the marsh hawk (Circus hudsonius) are common in migration and a few breed. Crows (Cor- vus hrachyrhynchos) are moderately numerous particularly in spring. In correlating these figures with the grouse population winter densities are most comparable. For the years 1931-32 and 1933-34, when the above data for resident predators were obtained, the mean winter density of grouse was 28 per square mile*. The corresponding figure for the total period of study was the same although it has varied from 18 to 10 per square mile. To summarize it may be estimated that the ratio of grouse to their principal predators in this area has averaged roughly as follows during the winter*^: grouse 28 per square mile fox 4 per square mile weasel 10 per square mile skunk 8 per square mile raccoon 4 per square mile great horned owl 5 per square mile Yet one must remember that these densities were by no means uniform over the tract and that minor changes from year to year have constantly taken place. Obviously variations in this pattern in other localities over the State arc endless but spe- cific data are largely lacking. Observations on the Adirondack area, however, indicate greater populations of weasels and barred owls; about the same of foxes* and raccoons: and smaller numbers of skunks, while great horned owls have not been obserxed. Cooper's and sharp- shinned hawks are slightly more numerous there during the summer. Goshawks are more fre- quent and breed sparingly in the region. The most common hawk is the broad-wing which seems to be negligible as a grouse predator. Crows are much less abundant. Occasional bob- cats (Lynx rufus) occur. Winter grouse densities on this area have averaged 31 birds per square mile, ranging from 17 to 39 per square mile in different years, .^t first glance this appears very similar to con- ditions on Coniiccticul Hill. Considering that oidy about half of the latter area is grouse cover, however, while all of liie other falls in this category, it becomes evident that the actual pop- ulation density was much lower in the continuous coverts of the northern region. VlILNERABII,ITY It is logical to sujipose that birds atid animals which are crippled or suffering from disease are more easily captured iiy prel>servalions at the Research Center indicate that * The trouifl cover on ihit area occupies roughly half the total acreage, A The akunk and rarroon hibernate For coniulerable pcriixls at ihii aeaaon. t All fnxea on this area were of the red speriro. t On the areas most intensively studied shooting has not been permitted. ROLE OF PREDATION 331 diseased birds usually retain their vigor up to a very short time prior to death. On the other hand an earlier decline usually takes place in cases of heavy parasitism. Physical fitness may also be affected by other influences. It has already been pointed out* that during the early brood period grouse chicks regularly suffer considerable losses from causes mainly other than predation. Undoubtedly the vulnerability of these birds is greatly increased at such times and predators frequently become the ultimate executioners. Then too there is some indication that certain combinations of weather conditions may lower the vi- tality of older chicks and thereby render them more easily caught. Similarly weather may decrease the resistance of adults during the late winter''. There has been no indication, how- ever, that starvation is of importance. Nevertheless, grouse, thoroughly up to par physically, become more susceptible to predation under some circumstances than others. An outstanding factor influencing such variations is the shelter value of the habitat. The availabiUty of escape cover is of the utmost importance in enabling birds to elude their natural enemies. Since in any covert there is a limit to the amount of such cover, surplus individuals become increasingly vulnerable as ihey are forced to occupy less favorable territories. This was well demonstrated in the instance already men- tioned^ in which one section of Connecticut Hill, after attaining the excessive fall density of a bird to 2.5 acres, still suffered a high loss the following winter. There is also evidence that during the winter birds of the year are more vulnerable than older individuals, both as a result of inexperience and because the better territories tend to be already occupied. At times snow-roosting birds may be more easily captured by certain predators, but at other times this habit may have just the opposite effect. Even under the best conditions the vulnerability of individual grouse is constantly chang- ing as they pursue their daily routine of activity. Among the chicks, particularly during the first few weeks of life, the relative "mothering" ability of the female undoubtedly is of great importance. Similarly, circumstances which cause a chick to peep increase its chances of being located and caught. Significance of Predation In the introduction to this chapter it has been pointed out that predation serves a very necessary function throughout the organic universe in helping to keep the numbers of all species within limits compatible with the environmental niches which they occupy. So it is with respect to grouse. Yet it may seem to the reader, particularly the sportsman, from the foregoing discussion of its effect in New York that predation is exerting an undue pressure on this game bird. A fact often lost sight of, however, is that adult grouse exhibit an intolerance of population densities above about one bird to four acres even in the best coverts. Like Daniel Boone, who it is said at one time moved farther into the wilderness when a family settled within some 20 miles of him, these birds tend to avoid crowding beyond that level. In other words, predators or no predators, surpluses over this saturation point may be expected to disperse of their own accord after the birds of the year have "come of age" in the fall. But most coverts are not of sufficiently high quality for this characteristic to be a regular influence. Yet this merely means that even lower limits are imposed in such cases depending on the nature of the cover*. Fall surpluses above this level tend to be eliminated by over- * See discussion of Role of Predation During Brood Period, p. 316. A See Chapter VI, p. 304. t See discussion of Predator Pressure, p. 327. t See Chapter XII, p. 523. 332 PRE DAT ION winter mortality of which predation has been the chief agent. Moreover, since there is an inherent maximuin to the carrying capacity of each covert the elimination of predation would only result in some other factor, possibly disease, sooner or later assuming its role. It can- not be emphasized too strongly that no data so far give any promise that the principle of compound interest can be applied to grouse or similar game populations. On the other hand predation during the nest and brood periods does represent a factor limiting the fall abundance which might otherwise occur. Under primeval conditions such a staggering of the total reduction which must take place is advantageous. Where man's pursuit of the bird for sport — being, as it is, confined to the fall — is important, however, the advan- tage is less real. But this point of view is valid only where such sporting activities are suf- ficient to utilize the bulk of any additional surplus it might be possible to produce. Unless so utilized they might better furnish dessert for some predator as eggs or chicks since few would survive to breed. Another function of predation is that of culling crippled, sick or otherwise defective stock. The Investigation has little direct evidence, however, of the importance of this service. In the first place crippled birds have not been present on the study areas since no shooting has been allowed. Furthermore, the incidence of disease has been negligible. Then, too, one marked female, in spite of a broken wing was known to have hatched her clutch of eggs and success- fully reared her brood. Nevertheless, such relationships may at times be highly valuable. Some effect of this kind probably occurs during the brood period since a large proportion of the losses at this time appear to result from causes other than predation. It is also likely that the effect of weather on adult mortality suggested in Chapter VI, if real, operates through fostering the decimation of the weaker individuals. Certainly there can be little doubt that over the period of its evolution the constant necessity of eluding its natural enemies has been a major influence in developing the very qualities which today make the grouse so prized a game bird. Finally, in considering its significance one must make a distinction between the fact of predation on individual grouse and the effect of predation on populations of this game bird. Obviously the inevitable reduction of its potential productivity must involve the death of a large number of individuals in one form or another, many by predators. But only under very unusual circumstances does such decimation endanger the maintenance of population levels in accordance with the carrying capacity of the habitat. Perhaps the greatest obstacle to an understanding of the role of this factor has been the failure to appreciate this difference. APPRAISAL OF THE VARIOUS SPECIES In addition to an understanding of the role of predation as a factor in the life equation of grouse the game manager must also know what predators occur in his locality and what their relative importance is. An appraisal of the various species in New York has been an integral part of this study. CHdisK Predators of the Northeast At one time or another <;rouse have probably fallen prey to nearly every carnivorous bird and animal occurring within its range, except those entirely incapable of attacking it. Be- yond this its eggs are often destroyed by a number of other species. Among wild mannnals the Investigation has recorded the red fox fVulpes fiilva), gray fox (Urocyon cinereoargenteus). New York weasel (Mustela noveboracensis), Bonaparte APPRAISAL OF THE VARIOUS SPECIES 333 weasel (M. cicognanii), skunk (Mephitis mephitis), raccoon (Procyon lotor), bobcat (Lynx rufus), opossum (Didelphis virginiana), woodchuck (Marmola monax), red squirrel (Sciurus hudsonicus), chipmunk (Tamias slriatusj, and mice. That other species, inhabiting the Northeast but not encountered in this work, must also be included in this category is indi- cated by the writings of other observers. Although citing no specific instance Eaton"' men- tions the mink (Mustela vison) and marten (Martes americana). Seton^ reports the lynx (Lynx canadensis) and fisher (Martes pennanti). The probability that wolves (Canis) take grouse where the two occur together is corroborated by the findings of Skinner"" in the West. Recently a case involving the porcupine (Erethizon dorsatum) has been published^. Un- doubtedly the eastern panther (Felis couguar) formerly included the grouse in its diet, while at the other extreme Allen'" has suggested that even shrews may menace young chicks. Where man's civilization has penetrated, the dog and house cat are added to the list. In this connection Ewbank'^ makes the interesting statement that in North Carolina razorback hogs destroy grouse nests. Among the birds have been identified the great horned owl (Bubo virginianus), barred owl (Strix varia), goshawk (Astur atricapiUus) , Cooper's hawk ( Accipter cooperi), sharp- shinned hawk (Accipter velox) and crow (Corvus brachyrhynchos). Then too a few instances have been observed in which pheasants ( Phasianus colchicus) have caused nesting failure by laying their eggs in grouse nests. Beyond these, various authors have noted a number of others. Thus Audubon"' reported the raven (Corvus coraxj. Forbush"* and Eaton"" the duck hawk (Falco peregrinus), McAtee''^ the red-tailed hawk ( Buleo borealis), marsh hawk (Circus hudsonius) and even one instance involving the broad-winged hawk (Buleo plalypterus), Hersey'"* the red-shouldered hawk (Buleo linealus), and Kipley"' the osprey (Pandion hali- aetus). Applying only by inference to the eastern race, Patton ''" states that grouse in the Black Hills of South Dakota are preyed upon by the Richardson's pigeon hawk (Falco co- lumbarius). Gross'" has added two records for the snowy owl fNyctea nyctea) and Sutton"' one for the screech owl (Otus asio). Undoubtedly several of the rarer species in this region such as the golden eagle (Aquila chrysaelos), gyrfalcons (Falco rusticolus) and the great gray owl (Scotiaptex nebulosa) should also be included but data are lacking and their signifi- cance is negligible. Finally a number of cases of nest robbing have been traced to the northern blacksnake (Coluber constrictor). Relationships During Each Life Period The foregoing is a rather imposing array. Yet only a few exert any appreciable influence on grouse populations, especially in sections where the latter are hunted extensively for sport. Furthermore, even among those species which are important, some are primarily effective in breaking up nests, others as predators of chicks and so on. The records of the Investigation indicate these relationships quite clearly for New York*. To what extent these data may be distorted by instances in which the victim was dead when found by the predator or "on its last legs" is often difficult to judge. Such a relationship certainly does not apply to nests. It probably occurs most frequently among chicks. With respect to adults, as has been discussed'^, circumstantial evidence by the process of elimination strongly indicates that in the majority of cases predation has been the original decimating agent. * The basis for diagnosing the species responsible for broken-up nests and birds found dead is described in the Appendix, p. 709. A See discussion of Role of Predation Durin[j Adult Period, p. 318. 334 PREDATION In interpreting the following data one must bear in iiiiiul that, in addition to reflecting the habits of the various predators, the picture is also colored by the distribution and abun- dance of these siiecies. The authors believe that the material presented is representative of the majority of the better grouse range in New York. At the same time it is quite probable that in other sections of the Northeast the situation may be entirely different. In fact in other localities species may be important which are not even mentioned here. Nest Period The degree of predation observed among grouse nests has been discussed under a previous topic*. For each nest so destroyed the species responsible was in most cases diagnosed. Table 40 summarizes these data. In this connection it should be pointed out that it is utterly impossible to secure an accurate picture of the egg-eating habits of a predator through stomach examination. TABLE to. PUOPORTION OF BROKEN-UP GROUSE NESTS ATTRIBUTED TO VARIOUS PREDATORS IN DIFFERENT REGIONS OF NEW YORK— 1929-1942 Species Adirondack CatskiU Rest of State Total Number Per cent Number Per cent Number Per cent Number Per cent ¥o% Weasel 20 .5 I 4 1 5 i i 2 i 43.5 10.9 8.7 8.7 2.2 10.9 '2!2 6.5 4.4 '2.2 37 10 io 10 5 43.5 11.8 11.8 11.8 5.9 1.2 * 12 — '2.3 113 34 36 47 32 9 8 9 3 <> 1 i 11 5 2 i 12 4 2 .34.1 10.2 10.8 14.2 9.6 2.7 2.4 2 7 o!9 0.6 0.3 0^3 3.3 1.3 0.6 (1.3 3.6 1.2 0.6 170 49 40 61 43 19 9 10 3 2 T 1 "» 2 1 15 4 2 .36.8 10.6 Fox or Weasel Skunk 8.6 13.2 9.3 Dog 4.1 1.9 Rpd Squirrel Red Squirrel or Chipmuok Chipmunk or Mice 2.2 0.7 0.4 0.2 Bobcat 0.2 House Cat ... 0.4 Crow 4.5 Great Horned Owl . . 1.7 0.4 0.2 3.2 0.8 0.4 Sub-total .... 46 12 100.0 85 21 100.0 332 55 100.0 463 88 100.0 Predator Undeterminable Total 58 106 387 551* * The total number of records used in this table differs from that indicated in table 35 because nests not comparable for iurrival calculations could nevertheless be included here. There is little question which are the outstanding predators in the State during this period. Over three-fourths of all the nests brokcn-iip have been attributable to the fox. weasel, skunk and raccoon. Following these, about tied for a [loor iiftii. arc the crow and dog. More- over their relative importance varies little in the different regions. By far the most serious is the fox. That thi> ajiplies to the red species just as much as to the gray has been amply attested by experience on Connecticut Hill where both are present and in the Adirondacks where only the red fox is found. Several times tliese animals have been observed in the act of breaking up a grouse nest. On one occasion such a drama was observed from a tree blind, \liont dawn one morning a red fox was seen approaching the vicinity of an incubating bird under observation. Finally it flushed her. This individual • Sre disruBHiim it( Rnlr of I*rc4lnti(in Diirinj; Ncm I'criml, p. 311. APPRAISAL OF THE VARIOUS SPECIES 335 gave every indication of associating the flushing of a grouse with the possibility of finding a clutch of eggs. After attempting unsuccessfully to catch her it returned and deliberately sniffed the base of one tree after another until it located the nest. In fact seven nests were believed to have been destroyed in one 200-acre wood lot by this same fox. Foxes have the habit of taking several eggs from a nest but leaving the remainder. Some- times they return, sometimes not. In fact, they have been known to leave a few even after the second visit. Losses of this type have been very small. Next comes the weasel, including both the New York and Bonaparte. Over most of the State the latter is the more common. Although it would appear from the table that the skunk might rank second, analyses of the available evidence indicates otherwise since a large proportion of the records listed as "fox or weasel" in actuality were the work of the latter. In no case did the evidence point to the mink but this certainly does not suggest that this animal would refrain from plundering a grouse nest if it found one. Regarding the hawks and owls they of course are not egg-eaters. Yet they now and then cause nest failure by killing the female bird. Similarly the pheasant sometimes lays its eggs in a grouse nest and when they hatch first the deceived hen sets out with the foster brood leaving her own to die. Occasionally a nest has been found when the eggs had all been removed and buried intact nearby, either under the debris of the woods door or down a hole in the ground. Apjiarently chipmunks or mice have been responsible. Similar activities of these rodents have been noted by King"° and Gross*. Brood Period As has been pointed out analysis of predation among grouse broods has been a most difiB- cult problem. Undoubtedly young chicks often fall prey to predators. But they are either devoured "hide, hair and all" or the remains are so quickly obliterated that the most intensive field work has failed to reveal them. A great many are thoroughly sound healthy birds. On the other hand there is considerable evidence that the bulk of these early losses is fundamentally the result of something other than predation^. TABLE 41. PREDATORS CONSIDEHKI) RESPONSIBLE FOR DEAD GROUSE CHICKS FOUND ON CONNECTICUT HILL AND ADIRONDACK STUDY AREAS— 1930-1942 Species Connecticut Hill Adirondack Total Number Per cent Number Per cent Number Per cent 1 15 '>2 ~1 12 2.0 29.4 43.1 2.0 23.5 1 1 6 i 1 7.7 7.7 45.1 30^8 7.7 2 1 21 22 1 16 1 3.1 Sharp-shiuned hawk Hawk ap 1.6 .32.8 34.3 1.6 Foi 25.0 Weasel 1.6 Total 51 100.0 13 100.0 64 100.0 Mortality during the latter two-thirds of the summer, however, seems more directly asso- ciated with this factor. At this time also remains begin to be found. In table 41 are pre- * GroBS. L. S., Circular Letter, U. S. Forest Service, Allegheny National Foresi, June 15, 1936. A See discussion of Role of Predation During Brood Period, p. 316. 336 PRE DAT I ON sented the diagnoses of the specimens which have been attributed to predation. Besides these, 21 other specimens were picked up to which no cause of death could be ascribed, but which it is probable were largely the result of predalion. In spite of the fact that only a small proportion of the chicks which are known to have died were ever found, it seems unlikely that many were taken by predators not included here. Furthermore, on both areas, the situation was similar year after year. It is fair to conclude, therefore, that these records are representative. The data indicate that the most important species have been certain birds of prey. While, except for a few cases, it has been impossible to determine the exact hawk or owl responsible, field experience leaves little doubt that the Coojier's hawk and sharp-shinned hawk are out- standing. Of the specimens referred to "hawk sp." nearly all were found on "killing logs" of these hawks, amid the remains of warblers, thrushes and other woodland birds upon which they regularly feed. In the group referred to "hawk or owl" — all from Connecticut Hill — the great horned owl undoubtedly entered in. A considerable immber of chicks were also taken by the fox, but whether this animal or the owl should rank higher it is impossible to say. As in the case of nests both the red and the gray fox appear equally destructive. The one record of a crow carrying off a young chick appears to have been very unusual. No definite evidence of predation on grouse chicks by other s|)ecies has been observed. To be sure it has occurred to a minor degree and under different circumstances might be im- portant. Certainly there can be little question that the goshawk takes a toll at this season when the two occupy the same habitat. The raccoon, too, will readily kill prouse if given a suitable opportunity, as was demonstrated at the Research Center when several females and their broods being held in natural enclosures were completely wiped out by this animal. The same may be said for ihe weasel. While losses from house cats have not been encountered this animal is undoubtedly a factor where it is more frequent in grouse habitats. On the Connecticut Hill study area in particular the red-tailed hawk, red-shouldered hawk and marsh hawk have been common. Yet no instances of their molesting grouse have been recorded during the Investigation. As their food habits sihow* their principal diet consists of mice and other forms chiefly found in open land types. Quite probably they occasionally secure a grouse chick, as for example the red-shouldered hawk examined by Hersey"". But in any event one may conclude that the effect of these species on brood survival is negli- gible. Similarly the broad-winged hawk, although the commonest one on the Adirondack area, was never associated with grouse mortality. Adult Period During the adult period a considerable loss from predation is customary each year, taking place chiefly in late winter and spring. The remains of a much larger proportion of these birds have been subsequently picked up by the Investigation's field crews than in the case of chicks. Analysis of this material, while obviously far from precise, nevertheless affords valuable data on relative predator importance not otherwise obtainable. Of the 1,174 speci- mens examined 550 have been undeterminable as to cause of death. The proportion of such records, however, has been qiiilc similar throughout the study, both fnmi year to year and seasonally. Those it has been possible to analyze have, therefore, been considered represen- tative. The diagnoses of those ascribed to predation are presented in the following table. • See ditcuMinn of KimmI Hibili in New Yurk. p. 343. APPRAISAL OF THE VARIOUS SPECIES 337 TABLE 42. PREDATORS CONSIDERED RESPONSIBLE FOR DEAD ADULT GROUSE EXAMINED BY THE INVESTIGATION— 1930-1942 Species Connecticut Hill Adirondack Other Areas Total Number Per cent Number Per cent Number Per cent Number Per cent 58 9 5 1 52 2o:( ')b 15 6 1 13.0 2.0 1.1 0.2 ILT 45.5 21.5 3.4 1.4 0.2 i 5 6 1 I's.-i 6.3 31.2 37.5 6.3 22 6 '2 ii 57 24 1 17.6 4.8 ■ 1.6 8.8 45.6 1"».2 1.6 0.8 80 18 5 3 1 68 260 126 18 7 1 13.6 (fOslmwk 3.1 0.9 (ircat Horned Owl or (lOshawk . hooper's Hawk or Goshawk. . . . 0.5 0.2 11.6 44.3 Fox 21.5 Weasel 3.1 1.2 Skunk 0.2 Total 446 100.0 16 100.0 125 100.0 587 100.0 The predators which have been found responsible for the great majority of this niortahty are the great horned owl and fox, both red and gray. In New York the former seems by far the most effective since in actuality it undoubtedly accounted for at least half the dead birds which could not be diagnosed beyond "hawk or owl." On the Adirondack area, however, this owl was not observed during the period of study although this is a purely local condition. It is significant that under these circumstances the total mortality experienced was very similar to that on Connecticut Hill and that it was asso- ciated with a greater proportion taken by foxes*, Cooper's hawks and goshawks. Over the State the Cooper's hawk appears to rank next to the fox, followed by the weasel^. During an average year this hawk is the more important in New York, aside possibly from certain j)arts of the Adiroiidacks, since a few winter regularly throughout the central and southern part of the State. The goshawk, on the other hand, is an irregular winter visitant from the North. While there is ample evidence in jiublished accounts of the destructiveness of this species when it appears in numbers no such invasion has been witnessed during this study. A few individuals were recorded on Connecticut Hill for the winters from 1933 to 1987 and of the nine kills attributed to it four occurred in 1937. In view of this it is prob- able that the majority of the records referred only to "hawk sp." were the result of Cooper's liawk activity. The higher proportion of records attributed to the goshawk under Other Areas (table 42) is due to the fact that most of this material was collected at Pharsalia where it happened to be present during the winters involved. Other birds of prey have not been implicated except for the finding of grouse remains in the pellet of a barred owl picked up on the Adirondack area. They, of course, may have taken grouse now and then but there is nothing to indicate that such instances have been more than occasional. In this connection McAtee^" found grouse in three of 754 red-tailed iiawk stomachs, in four of 601 marsh hawks, and in one of 145 broad-winged hawks. Gross"' found grouse only once in the many stomachs of the snowy owl^ which he examined. * Only the red fox is present in this locality. A No ditfercntiation has been made between the two aperies. New York and Bonaparte. t The same author has also reported grouse remains in one pellet of this owl.^'- 338 ['RED AT ION One of the most surprising statements in the literature is that of Townsend"~ in which he reports a crow carrying a full-grown ruffed grouse. Certainly such a feat is rare indeed. In the experience of the Investigation mammals other than the fox and weasel have been of little consequence to adult grouse. In coverts where stray cats are numerous, however, their depredations may become more frequent. Forbush"' reported 46 birds of these species killed by cats but did not state the area or the period of time involved. As has been discussed previously* the vast majority of the adult mortality each year takes place between fall and the following breeding season. This pattern appears to be primarily associated with the existence of surplus birds in the habitat. It is significant that the sea- sonal distribution of kills by the various predatory species has followed this very closely. Net Effectiveness Throughout the Year From the foregoing data it is evident that the predators most destructive of grouse are not the same with respect to the different life periods. It is, therefore. ai)pro])riate to state the conclusions of the Investigation regarding the net effectiveness throughout the year of the more important species. Over the better grouse coverts of the State the fox is easily outstanding. Not only does it take a considerable number of adults and chicks but it also has been responsible for over a third of all the broken-up nests examined. Second is the great horned owl followed by the Cooper's hawk both of which influence survival during all life periods since they from time to time take nesting females. Next comes the weasel, another species capable of taking grouse the year around but which is most important with respect to nests. These are the paramount grouse predators in New York with the addition of the goshawk when it appears in numbers. Beyond this, in those localities in the Adirondacks where the great horned owl is absent, the latter and the Cooper's hawk take proportionately more. Regarding the sharp-shinned hawk and skunk it is diflicult to say whether the number of chicks taken by the former outweighs the eggs destroyed by the latter. The raccoon ranks somewhat lower largely because it is less numerous than the skunk in most sections. Finally may be mentioned the crow and dog. Food Habits in New York Important as it is to know the effect of a predator on the game species with which one is concerned, an intelligent management program must also take into consideration its other food habits. Only in this way can a well-rounded picture of its jjlace in the environment be secured. It must be borne in mind that in no instance does a game bird of the Northeast constitute the staple food of any ])redator. At the same time even the taking of small numbers may rep- resent a degree of predation suflicient to have an important effect on the game population. The two approaches are thus complimentary to each other. Too often this has been lost sight of. The conclusions of the Investigation with regard Id tlu' effect on grouse have been dis- cussed. It remains to summari/e briefly the relali\c pnipDrlions of this and other types of food which comprise the usual diet of the more imi)<)rlaiit grouse predators in New York as well as some of those commonly suspected of being. Data of (wo principal kinds have been collected. First the stomachs of specimens taken in various parts of the State have been secured from time to lime and examined. These have * Sec diacii»i<>n <>( Itolr •> 8.8 5.5 4.4 1.1 20.9 10.9 10.9 Hurred owl 20 4 6.3 37.5 37.5 6.3 25.0 18.7 6.3 Snowy owl . 6 2 75.0 25.0 fioslinwk H .! 36.4 9.1 9.1 18.2 .36.4 18.2 9.1 Cooper's lliiwk . 11 I'l 4.5 31.8 4.5 9.1 45.5 9.1 18.2 Sliarp-shinned 53 23 3.3 13.3 83.4 Red-tailed hawk 76 19 8.8 40.1 19.3 7.0 1.8 7.0 22.8 31.6 hawk 45 12 3.0 45.4 21.2 12.1 18.2 l.Vl 15.1 Marsh hawk .... 35 14 14.3 38.1 14.3 28.6 4.8 14.3 The f (illowinp; comments on these data are considered under each species separately. Fox Most of the stomachs were those of red foxes although 20 grays were secured. Supple- menting these a total of 4.269 droppings have been examined. Of these 3.0.54 have come from Connecticut Hill and 719 from other areas in the southern part of the state, while 496 are from the Adirondacks. In the latter material only the red fox is involved, whereas in the other both species are represented. In table 44 the analyses of these specimens is tabu- lated in the same way as for stomachs. There is little to choose between the basic food habits of the two species. It is apparent that rabbits and mice constitute their staple foods throughout the State at all seasons. In the first group the varying hare is paramount in the northern part of the State, while south- ward it gives way to the cottontail. Among the mice the field mouse is taken most frequently, followed by the deer mouse. Lesser numbers of the red-backed mouse and other species have * The Btomacli analyses were made largely by tlie author altlioiigh a series of fox material was done by William J. Hamilton. Jr., of Cornell University. Pellet and dropping examinations were made in part by the author and in part by John C. Jones, working in collaboration with the Division of Food Habits Resrarcli of llie Federal Fisli antl \i'ildlifp Srrvire. Thomas Smyth and Ralph Palmer also assisted in the latter. A See Appendix, p. 875. t In carh species a number of tlie stomachs received were empty. These were deducted in figuring percentages. 340 PRE DAT I ON TABLE 41. PROPORTION OF DIFFERENT TYPES OF FOOD IN FOX DROPPINGS EXAMINED BY THE INVESTIGATION— 1930-1942 Typo „f r.K,.l Connecticut Hill Other areas outside Adirondacks Adirondacks Total Number Per cent Number Per cent Number Per cent Number Per cent Rabbit 1.617 1,323 97 325 109 17 175 220 571 901 539 53.1 43.4 3.2 10.7 3.6 0.6 5.7 7.2 18.7 29.5 17.6 334 242 13 81 32 8 54 75 149 268 81 46.4 33.6 1.8 11.3 4.5 1.1 7.5 10.4 20.7 37.3 11.3 293 1.56 50 66 50 .34 136 173 35 59.1 31.4 10.1 13.3 10.1 6.8 27.4 34.9 7.1 2,344 1,721 160 472 191 25 229 .329 856 1.342 655 52.6 40.4 Shri-w 3.8 11.0 (Irousf 4.5 0.6 5.4 Other birds 7.7 20.0 Fruit .... 31.5 15.3 3.054 719 496 4.269 been found. The fox seems to have a definite aversion for shrews, although they have been fully as abundant as the woods mice at least. Other mammals eaten have been primarily the gray squirrel, red squirrel, chipmunk and woodchuck. Of interest is the occasional occurrence of other predators such as weasel, skunk, raccoon, opossum and house cat. Analysis of the dropping material brought out a significant correlation during the fall and winter of 1935-36 between the irruptive abundance of gray squirrels and the frequency with which they were eaten by this animal. The same was true to a lesser degree of red squirrels. There seems to be a tendency for rabbits to be taken less during the summer than at other seasons and for such species as the squirrels, woodchuck and small birds to be tJiken more. Grouse make up a comparatively small part of Reynard's diet, although they have been taken more often in the Adirondacks than elsewhere. Both adults and chicks have been re- corded but in only one instance was an egg shell identified. This empha.-iizes the inadequacy of food habits studies of this kind with respect to certain types of predation, since fo.xes have been found to be by far the greatest destroyers of grouse nests*. FOX DROPPING CONTAINING GRt)USE IK AT HER KKMAINS The low incidence of grouse in the diet of this animal as compared with that of small mammals has also been shown by a number of other studies. Hamilton"' reported three in 206 red fox stomachs mainly from New York and one in 66 specimens from New England. Similarly English and Heimett in Pennsylvania found it to have been taken by 3.7 per cent of red foxes"" and 3.4 per cent of the grays" which they examined. • Src fliacuttion of Appraitjil of Ific Vllrioui Species During ifie NeBl Period, p. 334. APPRAISAL OF THE VARIOUS SPECIES 341 Weasel Included in the material examined were stomachs of both the New York and Bonaparte weasels but the number is insufficient to warrant separate treatment. There seems to be very little difference in the food habits of the two species, however. Mice were by far the most frequently taken items followed by shrews which occurred more often than rabbits. Grouse remains were not found althoufih the records of the Investigation show this animal to be one of the important predators of this game bird, particularly during the nesting season. Skunk Although the number of specimens examined has not been large all seasons of the year are well represented in the group. Insects were found more frequently than other types of food. At the same time rabbits and mice were taken by over a quarter of the animals and probably constituted a greater bulk. Fruits also were taken in season. Of the 77 stomachs holding food, four (all from the winter and spring) contained grouse remains — a finding which it is difficult to interpret since the skunk has appeared to be only a negligible predator of adult birds. In any case this is not an important source of food. That insects, fruits and small rodents constitute the ])rincipal diet of this animal is also evident in the data of Lantz"" and Hamilton"*. Raccoon The 'coon too feeds to a large extent on small rodents although considerable quantities of insects, fruits and grain are consumed when available. Of the 31 stomachs containing food seven held rabbit. Shrews and red squirrels were also taken. Grouse occurred once — a chick taken in August — and other birds not at all. Mice were not found either although Hamil- ton"' identified them in 17.4 per cent of the specimens he examined from the State. Mink Only eight of the 12 mink received held food. In one was found the remains of a mouse and in three muskrat hair while five contained miscellaneous items including a crayfish. House Cat Over three-quarters of the cats examined had fed on small mammals. Mice had been taken most frequently followed by rabbits, shrews and the Norwav rat. Also identified was a weasel. Birds were found in 2^ per cent of the material. Grouse, however, were not represented. Great Horned Owl In addition to the 115 stomachs of this owl a total of 1,517 pellets have been gathered from the southern part of the State, primarily Connecticut Hill. The species has not been observed on the Adirondack study area. The analyses of this material are summarized in table 45 in the same manner as the fox droppings were handled. Like the fox, this species relies chiefly on rabbits and mice for its sustenance. The rela- tive importance of the various species is about the same although the wood mice, especially the deer mouse, rank somewhat higher. Shrews also are mcjre frequently eaten. Among the other mammals red squirrels and chipmunks have been taken oftenest during most years. In 1935-36. however, as with the fox. the greater availability of gray squirrels was reflected in the owl's diet. But even more interesting was a very much higher incidence of flying squirrels in this year suggesting that they too may have reached a peak of abun- .342 P RED AT I ON TABLE 1.5. I'HOPORTION OF DIFFERENT TYPES OF FOOD IN PELLETS OF THE GREAT HORNED OWL EXAMINED BY THE INVESTIGATION— 1930-1942 TyfH-.if f.»,,l Connecticut Hill Other Areas Total Number Per cent Number Per cent Number Per cent Rabbit 111 "is 2:( K') u 5:i 40.6 .17.6 ».7 17.6 4..'-. o..'; 1.8 6.<> 1.1 4.2 144 IS 20 2<» 8 2 10 59.8 .19.4 8..1 12.0 3..1 0.8 OB 1.7 0.8 4.1 777 574 111 254 66 9 25 98 16 6.1 S1.2 .17.8 8.6 16.7 4.5 0.6 1.6 6.5 1.1 4.2 1,276 241 1.517 dance at the same time. A miiiiher of skunks were recorded as well as weasels, an opossum, crows, two screech owl.s. a Cooper's hawk ami a sharj)-shiiinr(l hawk. Grouse have occurred in onlv about 5 per cent of the material examined, a proportion very similar to that noted for the fox. Both adults and chicks have been identified. PELLET OF GREAT HORNED OWL SHOWING GROUSE REMAINS AND SUSCEPTIBIL- ITY TO DISINTEGRATING OF PELLETS COMPOSED OF THIS SORT OF MATERIAL Small birds were found more frequently in the stomach material than in the pellets. They were taken more often in summer than at other seasons. Miscellaneous items have included a considerable number of garter snakes, frogs, salamanders and crayfish. Analysis of 983 great horned owl stomachs from Pennsylvania*" also showed grouse to have been taken only a relativcK few limes il.l per cent) while small mammals, mainly rodents, were the [irincipal bill of fare. Barred Owl None of the barred owl stomachs held grouse and its remains have been found in onlv APPRAISAL OF THE VARIOUS SPECIES 343 one of the many pellets examined. The principal foods of this owl are mice and shrews, the latter being eaten more frequenth than by any of the other predators observed. Small birds and insects were also taken to some extent. Fisher"" likewise found small mammals to be the chief food of this owl, although he re- ported that in 100 stomachs poultry and game amounted to 4.5 per cent, mainly "half- grown fowls which roost among trees and bushes". Snowy Owl The four specimens of this owl containing food are too few to indicate much regarding its food habits. Gross'"', however, has reported the contents of 284 stomachs, 135 of which were empty, collected in New England during the flight of 1926-27. He found over 75 per cent had fed on small mammals, especially the Norvvay rat. A wide variety of birds had also been eaten including one grouse. Goshawk Although the number of specimens secured was small, analysis of their contents further demonstrates the tendency of this hawk tf) prey heavily on birds. Grouse occurred in one of 11 stomachs, pheasants in two and small birds in four, while small mammals had been taken by five of the birds. Of all the ra|)tors of the Northeast the goshawk comes closest to meriting the name "partridge hawk". Reporting on 243 examinations McAtee°" states "Grouse, chiefly ruffed grouse, were di'lcrniiiii'd in 31 of the stomachs". Tie also found birds of one kind or another in over lialf of the same group. Similarly in 538 stomachs from I'eimsylvania 153 or 28.5 per cent contained grouse and many others the remains of smaller birds, while about half held the remains of small mammals, chiefly rabbits and squirrels"^. Cooper's Hawk Like the goshawk the Coo])er's hawk feeds extensively on other birds. It preys largely on non-game species because they are more abundant and more within its capabilities. Yet grouse are taken just as readily whenever a suitable opportunity presents itself. Mammals had been eaten by onlv a little over a third of the s|)ecimens examined. Its preference for birds has been corroborated many times by other workers. Of interest is the high proportion — nearly half — of the stomachs of this hawk which have been empty. The same is also true of the next species, the sharp-shin. It suggests that their rate of digestion ma\ be quite rapid. Sharp- shinned Hawk This hawk preys even more predominantly on birds than either of the two preceding. Due to its small size, however, game species are seldom taken except as chicks. In the specimens examined b\ the Investigation over 80 per cent of its food has been small birds, the balance being rodents, chiefly mice. McAtee"' has reported a small variety of other foods. The records reported bv May"' follow the same pattern. Red-tailed Hawk This hawk, so frequentlv seen wheeling in broad circles over field and pasture, is pri- marily a mouser. Nearlv half of those examined had fed on these rodents, mainly the field mouse, while shrews, ralibits and squirrels were also taken in considerable numbers. 344 PRE DAT I ON Over 20 per cent had eaten insects. Among the other items identified garter snakes have been found in a number of instances. While no grouse were recorded one specimen contained pheasant remains. McAtee*" however, found grouse in three of 754 stomachs and it was also found in two of 193 rec- ords from Pennsylvania^'. Yet game birds comprise only a small proportion of its usual food. Red-shouldered Hawk Although no less a mouser this species — another of the soaring hawks — has taken small birds more frequently than the red-tail. No game birds of any kind were recorded in the food material examined. One instance of its feeding on grouse has been reported"*. McAtee'" has also noted a "strong taste for amphibians and reptiles"'. Marsh Hawk Beating back and forth over meadow and marsh this harrier hawk feeds mainly on small mammals, mice and rabbits pred(jminating. It does, however, prey to some extent on young pheasants and smaller birds when they are available. Randair'' reports that in Pennsylvania juveniles of many small ground nesting species were the most important items of food during June and July. Confining its activities almost entirely to open areas it seldom encounters grouse. Nevertheless, the latter were found in four of 601 stomachs reported by McAtee'". It is evident from these brief resumes that, for the grouse predators of the Northeast as a group, small manunals. mainly rodents, are the staff of life. Only in the "blue-darter" hawks do we find s|)e(ies whose principal diet consists of birds and among them only the goshawk customarily takes grouse as an appreciable proportion of its food. At the same time all predators are opportunists and quickiv take ad\antage of variations in prev avail- abilit) . I'hus deviations from the general rule may take place in accordance with local conditions. Interpredator Relationships Predators are usually thought of as spending most of their time harrying and killing the unfortunate species upon which they depend for food. One seldom stops to think that they also prey upon each other. Yet such strife among themselves goes on constantly in any habitat. The peculiar taste of the great horned owl for skunks has been noted many times. Weasels are also taken frequentiv by this owl and it has also been known to feed on many of the smaller owls as well as the Cooper's hawk, sharp-siiinned hawk and opos- sum. Likewise the fox now and then catches a weasel. Skunks, raccoons and house cats sometimes fall prey to this animal too. On the other hand, in the North, the lynx is a tra- ditional enemy of the fox. Very often young hawks disappear from the nest long before they are ready to fly. An instance of a red-tailed hawk feeding nn a freshly killed red- shouldered hawk has also been reported"'. To just what extent, however, such habits may serve to limit the numbers of the various species is still a problem for the future. There is also some indication of a reciprocal relationship between predator populations. This may be the reason why in certain areas the great horned owl is predominant, while in others the barred owl is most numerous. But data of this kind are few and fragmentary. THE PROBLEM OF CONTROL 345 THE PROBLEM OF CONTROL Early in this chapter it was stated that if, through predator control, the game manager could produce a greater fall surplus of grouse and the hunter be enabled to harvest it better sport would result. Whether or not measures of this kind will have such an effect can only be determined by actual trial in the field. The Investigation has carried on a few experiments of this nature. Experiments Conducted The first series of tests were made on land being surveyed in connection with the Con- necticut Hill study area. The tract to be used was divided roughly in half, 1,425 acres of grouse cover to 1,223 acres. Beginning October 1, 1930. an attempt was made to com- pletely eliminate all predators from the former portion although this goal was never attained. The other portion was used as a check. Nevertheless, between that date and August 31. 1932. a total of 557 individuals was taken*. Following this Nature was allowed to take its course for a year. Then between October 1, 1933 and April 30, 1934 elimination of predators was again attempted. This time the other jjortion of the tract was subjected to coiitri)! since there was a possibility the results obtained previously were unduly influenced by j)cruiiaritiei of the coverts chosen. During tliis |)criod 195 spciimens were removed*. Because the inilial experiment had been most effective in reducing nest losses and because foxes and weasels had been found the most important nest predators, the selective control of these two species oidy was a logical stej). This was first undertaken during the winter and spring of 1933-34 on a nearby area known locally as Hull Hill. The results, however, were inconclusive since even these animals were not successfully eliminated. Accordingly this se- lective practice was repeated on a portion (792 acres) of the test tract on Connecticut Hill from October 1, 1934 to April 30, 1935. During this time 18 red foxes, four gray foxes and two weasels were taken. Beyond this the fall of 1934 on this area witnessed the highest grouse density recorded by the Investigation — an average of a bird to 2.5 acres over a 187-acre covert. Even though this was a ])art of the area on which long-time population trends were being fol- lowed it seemed too good an opportunity to pass up. Intensive control of all predators was therefore instituted to determine to what extent such a level of abundance could be main- tained. From ()( lober 1. 1934 to April 30, 1935 a total of 97 predatory birds and mammals were renu)\c(l from this covert*. This completed the trials carried out direclb by the Investigation. Throughout all of them the usual grouse survey field work was carried on at all seasons on both the control and check portions. In 1940 a .separate project to study this problem was established within the Bureau of Game under the provisions of the Federal Pittman-Robertson Wildlife Restoration Act. As a study area. Valcour Island of 1050 acres, lying about a mile from shore in Lake Cham- plain, was chosen. A low population of grouse was present here as well as varying hares and cottontail rabbits. Complete elimination of predators was instituted and its effect on grouse and other species followed carefully. * A summary by species is inclu 19;U-l'«.-> Ycnr Per cent mortality CoiiLralled area Check area 19.31 21.4 39.0 20.0 23.0 51.3 1932 72.0 1931 19:i5* 37.1 *Cnntrol limitflH to foxes niiH wennels. It is at once apparent ibal. while llic general l<\cl (if iiiiirlalil\ varied ((insiderablv during these years, the cotilrollcd ;iica snlTcicd a inaikiilU Inuir loss each year llian llir clicik area. As discussed elsewhere practicalK all such losses ha\e been attributable hi predators. The difference, thcicfore, may be considered a dire'M the areas used were reversed as to treatment from the arrangement <>f 19.SI and 19.'^2. * A summary liy ■)>rrjri is incluJvU in itir Appcntlix. |i. 875. THE PROBLEM OF CONTROL 347 Another highly significant feature is the fact that in 1935 very similar results followed the control of foxes and weasels only, thus verifying the conclusion that they are the paramount nest destroyers in this region. The one covert where complete elimination of all predators was attempted in 1934-35 had too few nests to merit comparison although none of these were broken up. Similarly the number which have been located on the Valcour Island area have been too small to be indica- tive. That it has undoubtedly been effective on the latter area may be inferred from the fact that in 1940 only three broods were found (no spring census was made) and in 1941, after only six months of control work, three broods resulted from 13 breeders, while in 1942 a spring population estimated at 22 jiroduced 1 1 broods and 33 breeders produced 13 broods in 1943. Brood Period In contrast with the nest period the ])redator control carried on had little effect on losses among grouse chicks (table 471. As discussed under another topic* mortality during the brood ])eriod seems in large measure due to something other than predation. Beyond this the Cooper's and sharp-shinned hawks were, perhaps, the least well controlli'd of the impor- tant species on Connecticut Hill. Even on the unit where complete control was undertaken during 1934-35 a brood mortality of 73.9 per cent was recorded. T\Hr>E 17. COMPAHISON OF MHOOI) MOItTM.ITV ON \HK\ SrH-IKCTHD TO PHKOVTOR CONTllUL AND O.N CllliCK AKli.\— CO.NNKCTICLT HILL -1931-193.) Year Per cent mortality Controlled area Check area 1931 67.8 53.0 52.9 88.1 74.5 W.VZ 54.8 1934 51.4 1935* 79.7 ^Control limited to foxes and weaacls. On Valcour Island the mortality of three broods in 1940 before any control had been prac- ticed was 74.3 per cent. The next summer afler some eight months of work a loss of only 42.9 per cent was recorded, also based on onh three broods. That this cannot be construed as indicative, however, is demonstrated by the fact that in 1942 the proportion was 65.4 per cent in spite of the evident reduction of nesting losses as already noted and in 1943 it was 56.1 per cent. Adult Period Results at this time have been variable. The great majority of adult losses take place between fall and the following breeding season and predation is an important decimating agent. Yet control of predators has had no consistent effect on the degree of mortality. In fact, paradoxical as it may seem, in two of the four years it has been higher on the controlled area than on the check (table 48). Furthermore, the complete control attempted in one covert in 1934-35 failed to prevent a loss of 71.3 per cent, although some of this is attributable to dispersal. * Si'f discuaaions tif Kitlc of Pit-ilaliun During Briiod Periinl. p. ,^l(i. 348 P RED AT I ON TABLK 18. COMPARISON OF ADULT MORTALITY ON AREA SUBJECTED TO PREDATOR CONTROL AND ON CHECK AREA— CONNECTICUT HILL-IOIU-IQSS Year Per cent mortality Controlled area Check area 1930-31 1931-32 1933-34* 193t-35i 10.9 25.0 21.9 55.0 14.5 20.8 29.2 48.8 ♦Different method of computation used, see text. ^Cotktrul liiiiitnl to foxes and weasels. The figures given in this table represent the differential from September to September of the years indicated. In computing them complete census data have been available except for the fall of 1933 when only a portion of the area was surveyed. It has been necessary, therefore, to use some other means of evaluating the populations at this time on the two sub- areas. Since the whole tract was surveyed during the winter and sprinp census estimates were present for May 31, 1934. By adding to these the known mortality in dead birds actually picked u|) conservative figures for the preceding fall have been secured. While the ratio between these is considered representative it must be remembered that lioth are low with respect to the actual degree of loss which took place. It is interesting to consider these data. If predation is, as it appears to be. the major cause of mortality at this season, then it would seem logical that control of predators should increase survival. Yet the trials conducted have not had such an effect. Certain relation- ships have been noted, however, which may account, in part at least, for the results. In no instance has the differential between losses on the trial and on the check portions been great regardless of which was higher. Even in 1933-31 llu- difference cannot be considered important since, as pointed out above, the actual mortality in each case was some 20 per cent more. It is sigiiifuant, too, that this similarity extends to 1934-35 when only foxes and weasels were controlled. On Connecticut Hill the great horned owl has been the outstanding predator of adult grouse. In 1934-3.S it was not affected except in the one covert where com|)lete control was under- taken. A possible inference from this is that the operations carried on during the other years were unsuccessful with respect to this species, allowing il Ici niainlaiM its domination of the situation. That the number of these owls taken may not have acconiplished the desired reduc- tion invoKcs the fact that the great majority of those present on this area during the winter are transients. Under such conditions replacements would constant!) lend to offset the effect of control. Another influence which may have been involved stems from the fact that the two plots were adjacent as well as being surrounded by grouse coverts. Thus the "spring shuffle"* each yar would liml In have equalized both populaticms. Data from the unit on which a fall density of a grouse to 2.5 acres of cover was reached in 1934 indicate several things. In the first place dispersal to surrounding coverts as a result of the bird's intolerance of crowding took place innnediatel\. Beyond this, in spite of an intensive attempt to completely eliminate predators iri(li\ iclual> from adjacent territory quickly * See Chapter V, |i. 256. THE PROBLEM OF CONTROL 349 filled up many of the temporary vacuums created. In consequence a mortality insued even higher than that on the check area where, however, the initial density was less. With respect to Valcour Island losses in 1940-41 and 1941-42 were 23.5 per cent of 17 birds and 36.4 per cent of 33 birds. These mean little since the population densities were very low. On the Adirondack Area where conditions were comparable except that control has not been practiced a similar density in 1935-36 was followed by a similar degree of mortality. But the population on the Island, which reached 87 in the fall of 1943, suffered a mortality* the following winter of 82.7 per cent returning it to approximately its 1940 level. This oc- curred in spite of continued predator control. Thus it appears that predator control, especially on small areas, cannot be expected to maintain fall surpluses. Effect on Productivity In game management as in other enterprises "the proof of the pudding is the eating". The measure of predator control lies largely in its effect on the resultant survival of grouse. The experiments conducted have been effective in reducing nest mortality but have not appreciably lowered either brood or adult losses. Nevertheless, the increased nest survival on the control areas should have produced additional increments to the fall population. Unfor- tunately, the results arc inconclusive regarding this phase of the problem. They do, however, permit certain deductions to be made. It would seem logical that a comparison of fall population densities would be a suitable index upon which to base conclusions. Yet analysis of the data indicates that this is not the case. The fundamental difficulty lies in the fact that for various reasons the reduction in nesting losses accomplished on the controlled area did not consistently result in a higher den- sity of grouse chicks immediately following hatching on this area than on the check area (table 49). Thus, since mortality during the brood period, whether high or low, was not particularly altered by predator control, such a comparison is not representative. TABLE 49. RELATIONSHIP OF DENSITY OF GUGUSE BREEDERS TO TH\T OF CHICKS AT lL\T(.IIINr, AND TO THAT OF TOTAL POPULATION IN F\I,L ON CONTROLLED AND CHECK AREAS DURING PREDATOR CONTROL EXPERIMENTS ON CONNECTICUT HIL^-1931-1935 Population density (birds per 100 acres) Year Controlled area Check area Breeders Chicks al hatching Total fall population Breeders Chicks at hatching Total fall population 1931 . 5.0 8.7 7.3 7.2 19.6 22.4 21.6 17.5 11.4 19.0 16.3 8.8 5.9 8.3 7.2 10.2 15.7 22.5 20.6 21.1 9.8 1932* 18.4 1934 A 17.5 1935 12.2 •142 acres of controlled area omitted due to extraneous circumstances. A311 acres of check area omitted due to extraneous influences. The reasons why greater nest survival on the test area was not regularly attended by higher * Circumslanlial evidt-nre of disease was found but its significaii could neither be proved nor disproved. 350 P RED AT I ON densities of chicks as compared with the check unit are not entirely clear. A few, however, may be suggested. In 1931. for example, the density of the breeding population on the check area was higher than on the other, thereby tending to offset the greater nest loss. In 1932 there was some indication that renesting accomplished a similar result. .Again in 1935 grouse were considerably more plentiful in the spring on the check area which may he traced to the lack of effectiveness of control in preventing a high adult mortality on the test |)l()t during the preceding winter. But even densities at hatching time are not indicative because during two of the four vears brood losses were higher on the controlled area than on the check area. In 1931 a relatively larger number of chicks on the managed tract suffered a lower mortality and produced a greater abundance in the fall. In 1935 the reverse was true on the unit where selective con- trol was practiced. In 1932 and 1934 both June and September densities as well as brood losses were very nearly the same on both areas. It will be noted that the greatest apparent effectiveness occurred at a time when the general population le\el was low and rising. The suggestion has been made that lower brood densities may be subje inadequate for that ])urp()se. On the other hand there is a strong probabilit\ that the resultant fall density on the managed tracts themselves was greater than it would have been otherwise. .Assuming the same brood mortality would have taken place anyway, the increased nest survival could have no other result. And. after all. that is what the game manager is interested in. Turning to the question of progressively increasing the breeding population through pre- dator control the data are more clear. Outstanding is the record* of the covert which held an average of a grouse to 2.5 acres in the fall of 1934. Incidentally, tliis high level was reached in spite of a normal number of predators. On this unit, as well as on others in sub- sequent years, the grouse soon reduced their own density 1>\ (lis|>crsing a portion of the sur- plus'^ to less well populated surrounding coverts. Following this, in spile of efforts to ram- plelely control predators, the coincident influx of these species was suniciciit In cause a high overwinter loss just the same. Neither were these practices successful in the olln-r trials undertaken, particularly that on \ ali nur Island. True, if complete elimination of prcdation were actualK accomplished. sur\i\al unnid at first be greatly increased. That it would last long before being limited in some other way is highly improbabli'. Furthermore, control of such an inlcrisilN would seldom, if ever, be feasible. Thus the principal value of this type of management seems largely restricted to the reduction of nest mortality and then only on areas where the hunting pressure is great enough to utilize any additional surpluses created. Furthermore, these data indicate that control of foxes and weasels alone can be effective for the latter |)urpose in habitats similar to those studied by the Investigation. * Spr iliHrllKHinn involve bir»i« of itir ypar mainlv. ^^p^^^^^fe^- ''"'*'^^-. -^%^ CHAPTER VIII REPRODUCTIVE CAPACITY OF THE SPECIES By Frank C. Edminster and Walter F. Chissey REPRODUCTIVE POTENTIAL EFFECT OF SEX RATIO PROPORTION OF NON-BREEDING BIRDS Males — Females EFFECT OF AGE ON BREEDING Longevity and Senility NUMBER OF EGGS LAID Yearly Variations — Variation in Difierent Regions of New York RENESTING Number of Eggs in Renests FERTILITY OF THE EGGS VIABILITY OF THE EGGS INBREEDING ^ SUMMARY Normally, nearly all female grouse breed each s|)rin2. In some years, evidence shows that up to 25 per cent of them may fail to breed in limited localities. Non-breeding by males is unimportant, (p. 359). Both sexes breed the first year when they are about ten months old. (p. 359). An occasional wild grouse will breed when five years old or more. In captivity many breed at this age and a few have reached six years and bred. (p. 359). No indication of the reproductive capacity of grouse being affected adversely by variations in sex ratio was observed. Among newly-hatched chicks the proportion of males to females is normally about equal but as summer wanes males tend to predominate among birds of the year (p. 355). 354 REPRODUCTIVE CAPACITY OF THE SPECIES The average clutch size in first nests was 11.5 eggs. Fluctuations in average clutch size may result from changes in the proportion of old to young birds in the breeding population. Grouse apparently lay fewer eggs the first year. I p. 3011. Grouse whose nests are destroyed while laying or very early during incubation are likely to atlcin])t a second nest, but if their nests are lirokcn uji late in inculiation. the\ rarelv make another attempt, .^ince most nest losses occur late in incubation, renests do not ordinarily contribute largeK to the increase of birds, (p. 3641. Renests have averaged about seven and one-half eggs. No yearly variation of importance has been observed. ( jt. 364). Egg infertility has varied from one to seven per cent in different years, averaging 2.6 per cent. For renests it has been somewhat higher, averaging 4.3 per cent. This is largely due to occasional renests that are mostly or entirelv infertile, i p. 3651. Embryo mortality has been consistently low. Only 1.9 per cent of the fertile eggs in first nests and 3.9 per cent of those in renests failed to hatch. The variations in different years have not been important. ( p. 365). Inbreeding probably does not occur to any great extent and when it does, it exerts no ob- servable detrimental influence. I p. 367). At some period in the evolution of bird life, the cxait lime and place of whicii is lost in the haze of antiquity, a new species developed that might have been recognized as the proto- type of the ruffed grouse. From the beginning, its survival and increase were, as with any new species in its formative stages, largely dependent upon its ability to adapt itself to the surroundings and to reproduce in numbers at least sufficient to offset its losses. Among the factors controlling the abundance of any wildlife species, the capacity to re- ])roduce is fundamental, 'ihe inherent species characteristics that determine this capacity have cvoKcd frnm the adjustment of the forces lending to increase its numbers with those tending to liniil them. So long as these forces remain in balance, tlie species survives, but if reproduction should fail at any time, the species would be threatened with extinction. A species like the ruffed grouse, that suffers large normal losses each year, must have a relatively high reproductive capacity. The number of eggs laid and the fertility and hatch- abilit\ of these eggs must be large enough to provide for a sufficient increment each year to absorb these losses if the population is to remain stable. The proportion of female birds that lay eggs, as well as that of tin- males fertilizing them, nmst likewise be high. Other altribulcs of reproduction are similaii\ inipoilanl. HKI'RODUCTIVE POTENTIAL In order to gi\e a gra|)hii |ii( lure of lln- polciilial pKnlui li\ il\ nf lliis specio it mav be |)ointed out that one pair of grouse and their progen\ would ninnber 33.614 at the end of only five years if sex ratios were equal, all the females laid the average number of eggs* * Sincr the numbrr of cgB* per female hai averagcil 11.9 over the Rett of Slate regioii, 12 baa been uaed in llii» calculaliun in oriirr tii avoid decimals. EFFECT OF SEX RATIO 355 and all sources of loss were removed. In actuality, however, this re]3roductive potential is progressively diminished by a host of influences which translate it into productivity and populations of mature birds. These are con- sidered in detail in chapter XII. Only the inherent factors of reproduction are covered here. The subject is opened with a brief discussion of the sex ratio. While the proportion of males to females is primarily important as a prelude to productivity and will be discussed fully under that topic, its inherent tendencies are a factor of reproductive capacity. EFFECT OF SEX RATIO Among newly-hatched grouse chicks the proportion of males to females is normally about equal. Laboratory examinations of 464 grouse chicks under three months old, revealed 51.5 per cent to be males and 48.5 per cent females. In younger age classes the balance was even more perfect. As the summer wanes there seems to be a consistent tendency to a slight preponderance of males among the birds of the year. This often results in a few more males than females in the whole population through the fall and winter. Then the balance changes in the spring as the males suffer a somewhat higher loss as a result of courtship activities. In later s|)ring and through the summer, the po|)ulalion of females among the older birds generally exceeds that of the males*. There is no indication in the data gathered tliat the capacity for reproduction has at any time been adversely affected by the sex ratio. It is still conceivable, however, especially in marginal coverts, that the ratio of a scattered population might he so unbalanced as to prevent sufficient breeding to niairitain ibc |i(i|iulali(in level. PROPORTION OF NON-BREEDI\G BIRDS The proportion of a population that fails to breed during the spring season lowers the reproductive potential by that niiK li. Failure to breed may result from inunaturity (although not in the grouse), phvsiological dilliculties, or lack of contact with birds of the other sex. The evidence on this factor for each sex will be taken up separately. Males Determination of the proportion of male grouse that breed is difiicult for it is imjiossible to obtain enough direct observations. Hence one must rely upon the results of the spring mating season as evidenced by the degree of fertility of the females. This is not entirely satisfactory since it is apparent that, being promiscuous, even if every female nested and all their eggs were fertile, it would still he possible that some male grouse failed to mate. The data indicate that no appreciable failure in breeding among male birds has occurred. Only three nests of all that have been observed have contained complete clutches of infer- tile eggs. Conceivably, these could have resulted from lack of contact or from inability of a male bird to mate. They more likely resulted from physiological deficiencies in the females. The captive birds at the New York State Research Center have shown the same evidence of promiscuity that ajipears to be the habit in the wild''. Five females placed in a pen with a single male all laid fertile eggs. * Sci! Chapter XU. p. .SI 1. A See Chapter V, p. 1107. ►J o o o CO z tf) z "lil en z o o S tn u u 1^ K ■«: H ^ J < ^ u <; t/i u 3 O u cr: > o O ^ o z cc o s u s o 3 z K u Q ■< < lb K o u m ■ W'' «^'' ■i EFFECT OF AGE ON BREEDING 359 In general there seems to be no hesitation on the part of either sex in accepting a different mate, provided each is in the mating phase of the breeding cycle. If males do occasionally fail to mate, it does not, ordinarily, very seriously affect the number of eggs fertilized. Females Perfect mating is much more important with the females than with males. Every failure means infertile eggs. A small decline in the number of successful grouse nests in some favorite covert may well mean the difference between good and poor hunting the following fall. Also some females apparently do not attempt to nest and lay eggs. An accurate evaluation of this factor in the field is difficult, even where the population of females is known, due to the difficulty of locating all of the nests on an area as well as the impossibility of recognizing all instances of renesting. However, a fair estimate of the proportion of females nesting is possible through interpretation of known figures. The number of unlocated successful nests has been determined by subtracting the number of located nests from the number of broods found in the summer. By assuming that the number of unlocated nests that were broken up is in the same proportion to the number of located broken-up nests as the corresponding figures for successful nests, the number of unlocated nests that were destroyed may be estimated. The total number of nesting females may then be computed by adding the number of successful and unsuccessful nests, after adjustment for the number of renests. Finally, comparison with the known female pop- ulation gives an estimate of the proportion failing to nest. The degree of non-nesting obtained by this method has varied from zero to over 25 per cent of all the females on the Connecticut Hill study area in different years, the figures for 1933. 193Sand 1936 being especially high. That this is usually a minor factor, however, is evident from the fact that during most of the other ten years studied all known females were accounted for as attempting to nest. What may have been responsible for a high incidence of breeding failure when it did occur could not be determined. In a broad way it may be said that such females are probably physiologically upset and unable to breed properly. At least there is no observation to the contrary. But what the underlying causes of such variations may be remains an enigma. It has been suggested that, on the study areas, human disturbance of the breeding birds may have been involved, but this influence has not varied greatly from year to year. Thus, while losses resulting from failure of females to breed may now and then significantly reduce productivity, they are usually one of the host of minor influences on the grouse crop which, by themselves, are relatively unimportant but, which when added together, may measurably affect the hunting season population. EFFECT OF AGE ON BREEDING The ruffed grouse is sexually mature early in the first spring following birth. In captivity at the Research Center, they have usually bred during the first year, a conclusion likewise confirmed for wild birds by observation. In some years, every female known to reside on certain areas studied has at least started a nest. Since the whole population would inevitably include some female birds born the preceding year, these cases support the conclusion. One wild female was known to continue nesting attempts at least through five years. This bird was nest-trapped and leg-banded in 1936. That year she laid ten eggs. The next year 360 REPRODUCTIVE CAPACITY OF THE SPECIES and again three years later she was likewise caught at her nest. In 1937 her clutch was 13 eggs and in 1940 it was 12. The larger size of her second and, if she nested each intervening year, fifth clutches as compared to the first one observed adds weight to the evidence that the grouse usually lays a smaller clutch on her first attempt than in subsequent years. At the Research Center there is some indication to show that grouse lay fewer eggs after their fourth vear. They seem to build up to a peak of production when three vears old and to fall off slightly thereafter. One 6-year-old bird, which had previously laid larger clutches, dropped to ten eggs in 1940. However, the wild bird noted in the preceding paragraph still laid a normal sized clutch when at least five years old. The Investigtalion does not have evidence to show whether or not the birds continue to lay beyond their sixth year. However, such an advanced breeding age is unimportant as a factor affecting the reproductive capacity of grouse population. In New York, at least, only a very small proportion live to the ripe old age of six years. Longevity and Senility In 1901, Edgar A. Mearns'" wrote: "Mr. Samuel Eldred of Wakefield. R. I., purchased in December, 1900, a ruffed grouse from Mr. Joshua T. Bradley who stated that the bird was one of several young grouse caught by him in 1883 and marked by clipping off their hind and inner claws. This bird bore evidence of extreme age." And well it might, for if this record were accurate, the bird must have lived to be 17 years old. More recent literature and the present records of the Investigation indicate that the average adult grouse in the wild meets death before it is three years old. This is almost invariably a violent death and. therefore, does not indicate the potential longevity of the species. As has been mentioned, one banded wild grouse attamed an age of at least five years. At the Research Center, 4-year-old birds are fairly numerous, while 5-year-olds are not unusual. Six-year-olds, however, are rare and no bird has yet exceeded this age. At five or even six years of age. most grouse in captivity still ajjpear vigorous and healthy. Occasionally they exhibit the effect of long confinement by acquiring feather injuries or scalped heads, but the males mate successfully and females lay good clutches nonetheless. Wild birds seldom attain such an age. and hence, one concludes that there is but slight chance that the infirmities characteristic of senility could appreciably affect grouse populations. NUMBER OF EGGS LAID In considering reproduction, the number of eggs that a female bird lavs is important. If it were to change appreciably at any time, the possibilities for the hunting season might well be similarly changed. The evidence gathered by the Investigation in New York during the past 1 1 years, how- ever, does not bring out auv appreciable changes in egg laving proclivities. On the basis of 1,473 grouse nests examined between 1931 and 1941. the mnuber of eggs found therein has remained relatively constant. The minor fluctuations that have occurred were not such as to have materially affected the population changes that followed. Table .50 summarizes these records. It has been divided into |)arts representing the three areas of the state, as in other portions of the book, with the exception that the Connecticut Hill data have been se|)araled from that of the ''Rest of State". This seemed advisable because of the larpe number of nests observed on this studv area. NUMBER OF EGGS LAID 361 A hen grouse potentially might lay many more eggs than she actually does. A single bird reared at the Research Center was found to have 177 ova of various sizes in her ovaries, all visible to the naked eye. More than 150 ova have similarly been counted in a wild female taken just prior to the breeding season. A grouse might thus conceivably lay hundreds of eggs in its lifetime, whereas actually it will lay but a few dozen at most. The maximum number of eggs laid in a single season appears to be about 19. The normal average first clutch is 11.5, with an average of 7.5 for the renest, in case the first attempt is unsuccessful. What causes a female grouse to stop laying eggs as the season progresses is not certain. It probably involves some fundamental physiological changes. As has already been mentioned* temperature may have some effect. Likewise, a study of the influence of light, both as to duration and intensity, on the reproductive period of grouse was carried on at the Research Center during 1935 and 1936, In the winter and spring small amounts of light above normal were found to stimulate reproduction: large amounts at first stimulate, then inhibit, this reac- tion" '". It is entirely possible that the increasing intensity and the longer hours of daylight which characterize the advance of spring at first encourage, then, within the physiological limits of the birds, delimit the period of reproduction. Loss of weight has been considered as a possible factor in cessation of egg laying. It seems logical that this drain on the bird's system might so exhaust her as to cause a condition when egg laying is physically no longer possible. The available evidence fails to support this concept, however. Most of the birds" weight-loss takes place in the late winter, during incubation and in the early brood period. This rate does not ai)prcciably increase during egg laying. Furthermore, at the Research Center, both the males, and the females that do not lay, also lose weight. This is also typical of males in the wild^. The very fact that many birds, after laying a full clutch of eggs, lay another group follow- ing destruction of their first clutch, is evidence enough that the bird is not limited at this time TABLE 50, AVERAGE NUMBER OF EGGS IN FIRST NESTS— ENTIRE STATE— 1931-1941 Re|ion Total ^>ar Adirondacks CatskiUs Connecticut Hill Rest of Stale Number clutches Total eggs Average Number clutches Total eggs Average Number clutches Total eggs Average Number clutches Total eggs Average Number clutches Total eggs Average 1931 38 408 10.7 30 330 11.0 26 293 11.3 72 863 12.0 166 1.894 11.4 1932 35 378 10.8 27 300 11.4 29 336 11.6 90 1.049 11.6 181 2,072 11.4 1033 38 405 10.5 03 1,073 11.6 28 323 11.5 88 1,043 11.9 247 2,844 11.5 l')34 35 11.7 78 015 11.7 31 330 10.9 35 420 12.0 147 1.700 11.6 1035 46 11.5 61 663 10.0 41 506 11.5 12 129 10.8 121 1,344 11.1 1036 35 11.7 43 495 11.5 36 419 11.6 22 242 11.0 104 1,101 11.5 1037 11 11.0 17 207 12.2 17 204 12.0 39 473 12.1 74 805 12.1 103!) 18 209 11.6 25 284 11.4 33 396 12.0 46 567 12.3 122 1.456 U.O 1030 41 454 11.1 16 200 12.5 38 411 10.8 41 485 11.8 136 1,550 11.4 1040 43 472 11.0 6 68 11.3 40 581 11.9 19 297 12.0 117 1.348 11.5 1941 12 134 11.2 8 83 11.0 28 335 12.0 10 1I9 11.9 58 676 11.7 Total 236 2,587 11.0 404 4.632 11.5 359 4,143 11.5 474 5.617 11,9 1.473 16.979 11.5 » See Chapter \ 1, p. 303. A See Chapter II. p. 92, 362 REPRODUCTIVE CAPACITY OF THE SPECIES in its egg laying by physical inability to continue. On tho other hand, the smaller number of eggs laid in renesting attempts does give evidence that thr liirrl rn;i\ then approach the limits of its physical ability. A definite conclusion as to the cause or causes that bring an end to egg laying is not pos- sible with our present knowledge but the evidence points to changes in light intensity as the underlying factor, just as it is the stimulant that starts the production of eggs earlier in the spring. Yearly Variations It has been noted that the changes in the number of eggs laid per female year by year have not been such as to cause material differences in the following autumn pojjulatiDns. An exam- ination of table 50 will reveal, however, that some variation did exist in different years. Comparing state-wide averages, the extreme variations were between 11.1 eggs in 1935 and 12.1 eggs in 1937. The maximum change from one year to another was the increase from 11.5 eggs in 1936 to 12.1 in 1937. One might expect greater yearly variations from the records of a more restricted area since state-wide averages may tend to reduce irregularities. It is informative, therefore, to examine table 51 wherein the figures have been derived by regions. TABLE 51. GREATEST VARIATION IN AVERAGE ANNUAL CLUTCH SIZE (IN EGGS) Part of state In 11 years Between successive years Adirondacks Cnlskills 1.2 1.6 1.5 1.2 (I9.13-.M) .8 (lO.iUS) Conuocticiit Hill.. Host of Sliitc 1.2 (19.18-39) 1.2 (19.'»4-3S) These gross variations are significant* but no consistent trends are discernible to explain them. The Investigation has no evidence to show marked changes from year to year in the egg laying of binls of llie same age. However, there seems to lie a definite tendency for birds in their first year to lay fewer eggs than older ones! Thus the general average number of eggs would be affected if the jiroportion of youngsters in the |)(>pulafion changed appreciably. This actually is the case" and thus offers a means by which the observed yearly differences in clutch size may have come about. These changes do not. however, seriously affect grouse populations nor explain the great changes that take place in grouse numbers. Variation in Different Regions of New York The figures for the axcragc numlier of eggs per clul( h for the different portions of the State are given in table .SO. The average for the Catskills and for the Connecticut Hill study area is the same as the state-wide average. 11.5 eggs. The Adirondack nests averaged one-half egg lower and those from "Rest of State" four-tenths higher than this. Most of the latter group were taken from the southern tier counties in range similar to Comiecticut Hill. If these two are lumped, the average for the dis<-ontie(ied covert range is 1 1.7 eggs per nest. The difference * By Annlvni* fil Varianrr. A See Chaplrr XII. p. 513. NUMBER OF EGGS LAID 363 between this figure and the general average is of little importance. The one comparison that seems to hold real significance is that of the Adirondacks with the Rest of the State. The .7 egg difference, 11.0 to 11.7, between the Adirondack nests and all others in the State, is considerable since it represents 6 per cent of the number of eggs laid. This difference is significant by statistical analysis since the mathematical probability of its being true is more than the 19 to 1 required. A possible explanation of the lower egg-laying average of Adirondack grouse may lie in a repressive influence of the environment on the birds' reproductive ability. For grouse the car- rying capacity of the extensive forests in northern New York is far below that of the discon- nected coverts through the rest of the state*. It is at least a possibility that the level of egg laying might thus be affected. It is improbable that the physical ability of the female to lay eggs is reduced in the Adiron- dack environment, even though winter weather conditions are more severe and habitat generally less suitable than farther south. It is more likely that the rapid change of seasons during the breeding period may result in a shortening of the time when grouse have the urge to lay. The late spring causes a delay in nesting of about a week behind the rest of the State but by the middle of May the seasons have become equalized. Thus, there may be a seasonal time factor operating to give these northern birds a smaller average clutch size. A TYPICAL CLUTCH OF GROUSE EGGS The average number of eggs laid by Catskill grouse lies between that of the Adirondacks and the area of disconnected coverts. Likewise, the character of the Catskill range is inter- mediate in that the coverts are in rather large blocks, more so than is the case with the discon- nected coverts area but not nearly as continuous as the Adirondack forest. There may be significance in these parallels. Summarized, it might be that the carrying capacity of different types of range is an indicator of the average number of eggs the birds will lay. Regional * See Clmpter XU, p. 525. 364 REPRODUCTIVE CAPACITY OF THE SPECIES differences in egg laying, while small, exist and probably result from effects of the environ- ment. RENESTING The persistency with which the breeding females of a species of bird will renest- — that is, lay another clutch of eggs after one clutch has been destroyed — may well have an important part in determining its population. The mourning dove, for example, makes up for the very small number of eggs laid (only two) by repeatedly renesting over again whenever broken up. The grouse is not so constituted, however. Its ability, or at least its willingness, to try again following an initial failure is very limited. The time at which the destruction of the first nest occurs seems to be the factor determining whether or not she will renest. If a hen grou-se loses her clutch during the laying period or a very few days thereafter, it is probable that she will continue laying another set in a second nest. In this case the egg-laying period is merely extended. Once the hen has incubated her eggs for several days, the additional yolks that have formed in the ovary begin to be absorbed. Once this process has started, it becomes a difficult matter for the bird to resume laying. Nests thus destroyed generally meant a broodless female for that year. This limited probability of renesting woul.-l be less important were it not that most grouse nest destruction by predators occurs during the latter part of the incubation period*. Thus, most destroyed grouse nests are not replaced. On Connecticut Hill, where upwards of 50 nests are lost each spring, it is unusual to find evidence of more than half a dozen renests. Those that are found correspond consistently with the number of nests destroyed during the laying period. Number of Eggs in Renests The size of renest clutches has varied widely but has averaged consistentlv about seven and one-half eggs for all parts of the State, (see table 52). The rather small number of cases observed in any single year makes impossible the finding of any yearly variations that might have occurred. TABLE 5 2. AVERAGE NUMBER OF EGGS IN RENESTS— ENTIRE STATE-1931-1941 Region * Total Year Adirondacks CotakilU Connecticut Hill Rest of SUte Numbor clutches Total eggs Average Number clutches Ic.tal "ggs Average Number clutches Total eggs AvoruK*' Number clutcho-8 Total eggs Average Number clutches Total eggs Averagt^ 1931 5 26 5.2 4 28 7.0 4 34 8.5 6 41 6.8 19 129 6.7 1932 5 38 7.6 2 10 5.0 4 27 6.7 9 57 7.4 20 142 7.1 1933 6 45 7.5 6 41 6.8 1 8 8.0 17 122 7.2 30 216 7.2 1934 0 0 0 2 J7 8.S •» 12 5.0 1 7 7.0 5 36 7.2 1935 0 0 0 1 8 8.0 1 7 7.0 0 0 0 •> 15 7.5 1936 0 0 0 1 9 9.0 1 7 7.0 1 9 9.0 3 25 8.3 l'J37 0 0 0 3 27 9.0 3 18 6.0 14 114 8.1 20 159 7.9 1438 2 16 8.0 <) 0 0 0 0 0 3 23 7.7 5 .39 7.8 1939 B 66 8.3 <) 0 0 2 15 7.5 5 41 8.2 15 122 8.1 19»0 14 109 7.8 1 6 6.0 3 22 7.3 2 17 8.5 20 154 1941 4 34 8.5 3 25 8.3 3 22 7.3 0 0 0 10 82 8.2 Total 44 334 7.6 23 171 7.4 24 172 7.2 58 4^tl 7.6 149 1.119 7.5 • See Chiplec VII, p. 311. FERTILITY OF THE EGGS 365 FERTILITY OF THE EGGS When a pair of healthy grouse that are each in breeding condition, mate at necessary inter- vals, the eggs will be almost all fertile. Infertility may result where the hen bird is physi- cally impaired, where mating is imperfect, or there has been no mating. The fertility record of the eggs in wild grouse nests observed are summarized in table 53 for those cases where all the eggs could be accounted for*. TABLE .53. INFERTILITY RATES IN FIRST NESTS AND RENESTS— ENTIRE^STATE —1931-1941 First nests Reoests Year Number Total Number Per cent Number Total Number Per cent clutches eggs infertile infertile clutches eggs infertile infertile 1931 47 567 17 3.0 10 75 4 5.3 1932 61 723 17 2.3 7 50 5 10.0 1933 81 939 17 1.2 14 99 0 0.0 1934 25 272 19 7.0 1 6 1 16.7 1935 23 263 7 2.7 •> 15 0 0.0 1936 28 317 3 1.0 1 i 2 28.6 1937 28 336 4 1.2 8 62 4 6.5 1938 68 792 14 1.8 2 14 0 0.0 1939 63 717 oo 3.1 10 80 2 2.5 1940 51 597 24 4.0 6 47 3 6.4 1941 31 365 7 1.9 5 39 0 0.0 Total S06 5,688 151 2.6 66 494 21 4.3 The consistently high fertility rates in both first and renests are remarkable, especially when one considers the complicities of the breeding cycle. The yearly variations have been consid- erable— from one per cent in 1936 to 7 per cent in 1934. No evidence of trends from year to year that might indicate a relationship to population changes are discernible. Likewise, in no year was the frequency high enough to constitute a major threat. The average rate of infertility for 506 first nests was 2.6 per cent while for 66 renests it was 4.3 per cent. The rate of infertility in renests, while proportionately much higher than for first nests, is still not of any great significance. The 1.6 per cent increase obviously is not a large factor in grouse production but it does indicate a tendency for late breeding to induce more infertility than is normal among first nests. There is, then, no evidence in this 11-year record that infertility is a very big factor in determining grouse numbers. VIABILITY OF EGGS The embryo of a fertile egg is a living organism that may at any time during its develop- ment period be subjected to adverse influences and, as a result, may die. Any one or more of several causes may bring death. Among the more common causes may be listed inadequate incubation, an inherited weakness in the germ, or some structural failure in the egg itself. The cause of death in any particular instance cannot be identified except in rare cases. The records for 462 first and 64 renests for which data are complete are summarized in table 54. * When the shell remaina plus the unhatched eggs failed to eqiuil itie [iiu>\vri rlulili nuinher. tlie record was not used in the tables on infertility and embryo mortality of the eggs. Records from wild eggs that were collected and placed in incubators at the Research Center were felt to be representative of fertility and are included in the tabic. 366 REPRODUCTIVE CAPACITY OF THE SPECIES TABLE 54. EMBRYO MORTALITY IN FIRST NESTS AND IN RENESTS— ENTIRE STATE— 1931-1941 N. Y. State except Connecticut Hill Connecticut Hill Year NumbtT clutchi*3 Total ejigs Total fertile eggs Number dead germs Per cent dead germs* Number clutches Total eggs Total fertile eggs Number dead germs Per rent dead germs* Firet nests 1931 38 52 75 11 2 IS 22 37 40 33 13 454 619 874 123 21 162 267 448 454 374 151 440 605 862 115 21 161 265 441 444 355 149 6 9 5 3 0 :t 8 10 0 3 2 1.4 1.5 .6 2.6 0.0 1.9 3.0 2.3 0.0 .8 1.3 9 9 6 14 21 13 6 15 10 9 12 113 104 65 14') 242 155 69 186 108 111 143 110 101 60 138 135 153 67 IHl 108 107 138 11 4 3 10 4 6 0 4 6 10.0 1932 4.0 1933 5.0 1934 7.2 1935 1.5 1936 1937 2.6 3.0 1938 1939 3.3 0.0 19*0 3.7 1941 . 4.3 Total 338 3,947 3,858 49 1.3 124 1.445 1,.399 52 3.7 Renesta 1931 8 7 14 ' 1 "s 2 8 5 2 58 50 99 ' 8 62 14 65 41 17 54 45 99 "a 58 14 63 38 17 2 4 1 0 "s 1 0 0 3.7 8.9 1.0 0.6 's.it 7.1 1.6 0.0 0.0 2 '2" 0 2 17 ' 6 7 7 'I's 0 14 17 ■ 5 7 5 1.1.' 14 3 0 0 0 i' 0' 17.6 1932 1933 1934 0.0 1935 1936 0.0 0.0 1937 1938 . 1939 1940 1941 6.7 0.6 Total 55 414 396 14 3.5 9 66 63 4 6.4 *Calculated in relation to total fertile eggs. Upon examination of the egg viability records for all sections of tlie State, it was at once apparent that only one group failed to follow the general trend. Connecticut Hill nests had a consistently higher loss and for this reason are listed separately in table 54. The explanation for this discrepancy in the Connecticut Hill nests apparently lies with the Investigation workers and not with the grouse. From the time when first located, these nests were observed once each week until hatched. In spite of special precautions taken, some of the birds were disturbed and flushed several times. This disturbance seems to have resulted in some additional inadequacy of incubation beyond that usually recorded. On the other hand, most of the nests found over the State were not disturbed after discovery. The final record was made after the hatching date had passed. Because of this disturbing factor, the state wide data, excluding Connecticut Hill, are more typical than the records as a whole. This group exhibits a notably narrow range of varia- tion in embryo mortality fron year to year. Among the first nests these losses ranged from zero in 1939 to 3 per cent in 1937. The 11-year average loss was 1.3 per cent. For renests, the upper extreme and average were both greater. The highest loss was 8.9 per cent, occurring in 1932. The number of observations of renests each year was rather low and hence the larger losses observed in the embryos may be somewhat exaggerated. However, the average loss of 3. .5 per cent of fertile eggs is derived from quite adequate data and, there- fore, some diHercnce appears valid. INBREEDING 367 The higher average losses in Connecticut Hill renests compared with all others must be discounted owing to the small number of observations involved. The differences in average embryo loss in first nests and in renests is considerable, being nearly three times as great in the second nests. However, it is still only an additional 2.2 per cent of all the fertile eggs and does not involve losses of importance. It must be concluded, likewise, that all lossss from embryonic death are of minor impor- tance as compared with many other causes of grouse mortality. A NEST FULL OF GROUSE CHICKS JUST HATCHED (SEVERAL SHELLS REMOVED BY PHOTOGRAPHER) INBREEDING A pet bug-a-boo commonly suggested in recent years, as a possible contributing cause of the grouse cycle, is inbreeding. It was first mentioned by Stoddart'" but has since received more and more attention in succeeding years. To many it seemed simple and logical. The sedentary habits of the bird in times of high populations was thought to result in breeding within families. This, in turn, was said to cause deterioration of the species and sharply reduced numbers. The concept does not stand up well in the light of numerous pertinent facts. In the first place, inbreeding is not necessarily detrimental to the offspring. Only when the parents possess serious genetically transmissible defects could inbreeding be a problem. As Leopold'™ points out, wild species are of relatively pure strain and the likelihood of serious consequence from inbreeding is much less than in domestic animals where little natural culling takes place. Inbreeding may operate to a species' benefit as well as detriment. It has proved useful in developing superior strains of some domestic animals. If it were to injure wild animal populations, one may assume that it would show up quickly in isolated populations. Here, the opportunity for inbreeding would be greatest. Such might be the case where birds were stocked in a habitat not previously occupied by 368 REPRODUCTIVE CAPACITY OF THE SPECIES grouse and where the succeeding; generations would all be derived from the stocked birds. Yet the one known case of this kind with grouse attests to the prolificacy and hardiness of large groups resulting from such plantings. In this instance, 19 grouse were released on Anlicosti Island in the Gulf of St. Lawrence. These increased until the species was fairly common a few years later. The population has since persisted in the face of a high predator population. Inbreeding is only likely to occur with birds that are strictly and wholly sedentary. Other- wise, blood strains are continually interchanged. As has already been noted, the grouse is by no means wholly sedentary. The young birds move about considerably prior to their first breeding season. A large proportion of all grouse select a breeding territory different from the place where they were raised. Since grouse are probably promiscuous in their breeding habits, the opportunity for inbreeding is further reduced. Thus one must conclude that inbreeding seldom occurs in grouse and that there is no indication of its being detrimental if it does. Thus a ghost is laid. •.■;y;;fg%iUi< CHAPTER IX INFLUENCE OF MAN By Frank C. Edminster AS A GROUSE HUNTER Number and Per Cent Bagged — The Hunter Take — Average Bag, Seasonal and Daily — Crippling Loss — Total Hunter Kill — Success PIatio — Effect on Grouse Abundance of Man's Hunting. AS A HUNTER AND TRAPPER OF PREDATORS Number and Per Cent Taken by Hunting — As a Trapper of Fur Species AS A FARMER Clearing of Land — Maintenance of Openings - — Pasturing of Livestock — Other Domestic Stock AS A LUMBERMAN AS A CONSERVATIONIST Laws — Commercialization of the Take — Seasons for Hunting — Bag Limits — Methods of Take — Predator Control — Refuges, Sanctuaries and Land Posting. EFFECT OF MAN'S AIDS ON GROUSE The Axe and Plow — Fire — Automobiu:s and Highways — Guns, Traps and Snares — Grouse Dogs <£< SUMMARY The total hunter kill in New York State was found to approximate 17 per cent of the pre- hunting season grouse population. This total kill may be divided roughly into five-sixths birds retrieved, or hunter "take", and one-sixth cripples, shot but not retrieved, (p. 373). The average seasonal bag of grouse per grouse hunter has varied from .95 to 4.9 birds per hunter in New York State, (p. 3751. The daily bag of grouse hunters on check areas was about one-third grouse per actual hunt- ing day of 5?4 hours. Only one in four was found to have taken one or more grouse in tlie average day's hunt. (p. 375). The normal success ratio of grouse hunters varied from 7 to 11 per cent of grouse flushed 370 INFLUENCE OF MAN that were actually bagged. The average hunter brought to bag about one out of every four birds shot at. (p. 378). The general effect of man's hunting on grouse, as currently practiced, is not detrimental and may even have a positive beneficial effect during years of high populations, (p. 379) . The number of grouse predators killed by hunters in New York is too small to exert an important influence on grouse, (p. 380). The trapping of predatory species for fur, with or without the stimulus of a bounty, is neg- ligible in its effect on grouse abundance, (p. 382). Farmers, in the Northeast, are of first importance in determining the condition of grouse habitat, (p. 383). Land clearing operations, where not too extensive, have improved the productivity of the range; where too extensive, they have exterminated the grouse from large areas of potential range. The maintenance of openings in the form of farm fields carried on by cropping and pasturing is fundamental in preserving a high grouse carrying capacity. (p. 383). Domestic stock, when pastured in woodland and overgrown areas, serve a useful purpose in maintaining desirable openings but in other instances are destructive to grouse habitat, (p. 384). Lumbermen have exerted both a favorable and unfavorable influence on the condition of the habitat on which the number of grouse largely depends, (p. 385). Proper conservation legislation is a prerequisite to good game management. Proper handling of the grouse harvest requires executive authority in the Conservation Department for the proper regulation of seasons, bag limits and other controls of the harvest, (p. 388). Man's tools have exerted a profound influence upon grouse. Primary among these are the axe and the plow. Guns, traps and snares were formerly used to take grouse in great numbers, (p. 392). Automobiles and highways have made distant grouse ranges accessible to the city dweller. On the other hand, the abandonment of many rural highways have made local areas less accessible with the more modern vehicles, (p. 394). Grouse dogs still play an important part in the day's hunt, although their numbers are less than a few decades ago. ( p. 395 ) . While man's direct effect ujioii grouse numbers does not often rank him as a limiting factor, still it inav be said that, within certain limits, the future of the grouse rests with him. Man not only has a direct inllueiicc on grouse numbers through his activities as a hunter but abo exerts numerous vitally important indirect influences. He is a many faceted factor. As a hunter, he not oidy kills grouse but affects them through killing their enemies. In a still more indirect sense, his activities serve to increase or decrease the jirimar)- foods of major grouse enemies, notably some of the rodents, and thereby affect the habits of the preda- ^5 A GROUSE HUNTER 371 tory animals themselves. As a farmer, he cultivates the land and harvests its crops. Through these operations he largely determines the nature of the cover and, hence, the carrying capacity of the land for grouse. In those areas where he does not farm, he often assumes the same role, in so far as grouse are concerned, as a lumberman. Probably most significant of all is the gradual evolution of man as a conservationist. In this role he modifies his actions in a deliberate effort at producing more of our renewable resources. By affecting the numbers of grouse directly and through controlling its habitat, it is clear that he can, within limits, increase or decrease the numbers of grouse according to his will. AS A GROUSE HUNTER The grouse, along with other game species, played an important part in furnishing food for the early settlers of the Northeast. But it was not until the beginning of the nineteenth cen- tury that many written references to grouse hunting were made. Then, it was largely a mat- ter of replenishing the home larder or of supplying the city markets. Wilson*" indicates the lack of "sport" in grouse hunting at this time when, speaking of the advantages of a good dog. he says, "the more noise he keeps up seems the more to confuse and stupifv them, so that they may be shot down, one by one, till the whole are killed". In fact, they were such easy prey that traps, snares and deadfalls were in more prevalent use to bag them than guns. Audubon's" recording of the sale of grouse in the Cincinnati market in 1820 for 12''/; cents each indicates that large numbers (in proportion to the demand) nnisf have been killed. Elliot'" relates of one firm that received 1.000 birds on each Saturday for manv weeks in suc- cession. Knowing the strict limitations of transportation in that era. it is evident that huge numbers must have been brought to bag. Market hunting flourished through the greater part of the 1800s. rising and falling with the abundance of the birds but succumbing gradually (as a legitimate business) to the restric- tions of new laws*. Monon"" indicates that this "business" died a hard death when he says, "the whole northern country, from Amsterdam to Northville. New York, is infested with pot-shooters who hunt (out of season) for the Saratoga market". Nevertheless, from this period on there was a gradual transition in grouse hunting from "pot-shooting", both legal and illegal, to sport. By the turn of the century the change was about complete except for a small amount of illegitimate market hunting which continues even today. One of the few records indicating concretely the quantity of birds killed is given by For- bush"° who quotes a Mr. E. F. Staples of Taunton, Massachusetts. This gentleman said that in the early 1880s about a thousand birds were killed in a season on the 20,000 acres that he ranged. While an estimate is not given for the total number of grouse on the area, he does intimate that they were "real good" years — the last good ones in his estimation up to time of interview (1908). With a density then equal to the peak density of the recent period of abundance, this would have meant about 5,000 grouse and a bag of around 20 per cent. As the twentieth century progressed, the interest in grouse hunting as a sport increased rapidly and concern over the periodic disappearances of the grouse grew each time thev became scarce. Volumes were written on the subject and over-hunting was one of numerous * See Chapter I, p, 17. 372 INFLUENCE OF MAN "explanations" prominently mentioned although no one attempted to actually find out what proportion of the grouse actually were taken by hunters. Consequently, among the first assignments of the Investigation was an evaluation of hunting in the field. \UMBER AND PeR CeNT BaGGED One measure of the legal harvest of grouse in New York is through the report required of sportsmen by the clerk upon issuing a new license. It is recognized that these reports are not entirely accurate, but the errors are probably the same one year as another. Hence, allowing for this, they are valuable in indicating trends. These are given in table 55. TABLE .55. REPORTED KILL OF GROUSE AND AVERAGE SEASONAL BAG PER LICENSED HUNTER REPORTING IN NEW YORK^1923-1939 Year Ij^ngth of hunting season Number of grouse reported killed by licensees Number of licensees* reporting game bagged Average number of grouse bagged per hunter reporting Average number of grouse bagged I>er grouse huntert -\verage Humbert of grouse per grouse hunter per month 1923.... IH months 1^ months 1>^ months IVi months 1 V-^ months no season no season I'a months 3 weeks 3 weeks 1 month 1 month 1 montli 1 month 1 month 1 month 1 month 134,541 94,797 99,431 118.720 80,680 '36,488 40,448 55,845 68,940 84,614 128.301 80.282 94.056 140,851 157,151 165.122 A 202,661 A 204,525 A 180.060 220,920 194.779 163.597 133.980 149,710 156.650 153.379 136.521 147.332 165.811 166,876 .81 .47 .49 .66 .37 .19 .25 .41 .46 .54 .84 .59 .64 .85 .98 4.05 2.35 2.45 3.3 1.85 !95 1.25 2.05 2.3 4^2 2.95 3.2 4.25 4.9 27 * 1924 1 57 1925 1 63 1926 22 1927 1 23 1928 1929 19.30 63 1931 1 67 1932 2 73 1933 2 3 1934 1935 4 2 1936 1937 2.95 32 1938 4.25 1939 49 *An average of 61 per cent of all license holders make a report, of which 54 per cent (or 33 per cent of all licensees) report tak- ing game. AOnly hunting licenses were issueil these years, of which it is assumed an average of 61 per rt'iit reported, as in other years in which combination hunting, fishing and trapping li<'enses were issued. tliased on assumption that 20 per cent of all iNew ^'ork licensed hunters rejMjrting their game bag are grouse hvinters. JAverngc number of grouse bagged per grouse liuiiler wlien the length of the hunting season is correlated to the basis of 1 month. In interpreting these figures, certain facts should be noted. Tiic bag limit.-i in all years were three per day and 15 per season. The figures are not weighted to account for licensees not reporting nor for hunters not licensed. Only part of the licensees reporting game bagged were grouse hunters. When allowances are made for these factors, estimates of the average number of birds taken each year bv the grouse hunter is approxiinatelv the same or slightly lower than experi- ence leads us to expect. While this annual grouse bag per hunter is small, it compares very favorably with that of other species. The total yearly kill is. no doubt, greater than re|)()rted since some licensees do not report, landowners do not need licenses to hunt on their own lands, and some illegal hunting is done. No satisfactory method of taking these factors into account is available, so any estimate of (he actual kill is at best only a guess. It is. probably, at least double that reported. There are several interesting facts to be gleaned from these figures. In direct confirma- lion of the opinion of many "old-timer.s", there have been more grouse in the current peak years than in several decades. It is very noticeaiilc that the average take per hunter has materially increased, while that for the estimated grouse iiuntcr has nearly doubled. Although these figures meet the general appraisals made on experience alone, those for the later years may be somewhat high since, with the increasing niiniln'rs of grouse more and more hunters are turning to them for their sport. AS A GROUSE HUNTER 373 During seasons of few grouse, a larger percentage of the hunters became pheasant minded and did not hunt the partridge. Therefore, our average figure of 20 per cent of all licensees reporting game taken being grouse hunters may be a little high. And. during abundance of these birds, with more gunners taking to the uplands, that average may be a trifle low. Even with these allowances, it is still apparent that each sportsman has had a better bag during recent years. All of which means that, during the past ten years in New York, grouse have been more than holding their own in spite of increasing hunting pressure. The Hunter Take Leaving trends, now let us turn to the specific records of the number of grouse bagged. The first field studies to determine the effect of hunting on the grouse population were made by the Investigation during the 19.30 and 19.31 hunting seasons. Check areas were chosen in Tompkins County in the hillv. south-central portion of the state. For the 1930 studv. eight areas were chosen, of which six proved satisfactory. In 1931. twenty-five areas were checked, of which thirteen proved acceptable. Censuses were taken before and after the hunting season and reports were obtained from the hunters using the areas. Table r>() sum- marizes the grouse kill records for both studies. TABLE 56. GROUSE KILLED BY HUNTEK.S ON HUNTINCJ C.HEC.k \lii;\S 1<):!(I-I'>:U Year Number of reports received Number of grouse on areas before hunting season Number of grouse killed Per cent bagged 1930 1931 20 79 62 186 6 29 9.7 15.6 In 1930, the open season extended from the first of October until the fifteenth of Novem- ber. The grouse population was at about 30 per cent of its peak. The 1931 open season extended from October 26 until IVovember 15 and the grouse population was about 60 per cent of peak abundance. The per cent iiagged is based on all coverts. In interpreting these figures, it must be recognized that in 1930. when the season was re-opened after two years of closure, a large share of the hunters did not approve of the open season and refrained from hunting grouse because they believed the birds were still too scarce. This reaction is reflected in the low rate of take for that year. In 1931. with a very definite continued increase in the grouse population in spite of the 1930 open hunting season, this feeling was less widespread and the results for this year were probably more nearly normal. Another hunting check was made in the fall of 1936 on 1.379 acres of public hunting grounds. In a season extending from October 21 through November 1.5. hunters bagged 13 grouse. These represented 13.9 per cent of the pre-season grouse population estimate of 94 birds. The highest take on any one of the coverts was 23 per cent. This study simply adds weight to the earlier conclusions, since the results were very nearly the same. Following the first New York studies, two other similar hunter-check surveys were made in the mid-west. \^1iile each involves different ecological conditions than New York, both are valuable for comparison. Saunders* working in Michigan found the hunter take in 1932 to be 6.7 per cent of the September population. This figure is no doubt lower than the actual take since the pre- * Rulil. H. D.. personal letter lo the author. 1936. 374 INFLUENCE OF MAN season census was taken very early. The bag in 1935*" on one area, Pigeon River State Forest, was 17.3 per cent while on a second area, Escanaha River Tract, it was 17.4 per cent. But the corresponding figure — percentage of September population — taken on the same area as in 1932 proved to be only 10 per cent. Thus the actual hunter take in 1932 was probably nearly double that indicated. The results correspond remarkably close to those secured in New York. Trippensee's analysis of three areas in the Superior National Forest, Minnesota, on which both hunter tallies and before-and-after hunting season censuses* were taken, vary widely as to hunting take. In one area of 1,280 acres the take was only 9.4 per cent whereas in another of 2,560 acres, it was 38.2 per cent. The average for the 6,400 acres in the three areas was 19.7 per cent. While apparently more variable, these figures are but slightly higher than those of Michigan and New York. ■ ^ ^Bok'jfl PARTRIDGE -. ■ HELP MINTED K" \ ^^H y 64. hI ^^^^^^Vj EXPLANATORY POSTERS AND REPORT CARDS WERE TESTED AS A METHOD OF SEClfRING HUNTER KH.I, RECORDS While not based on direct field studies, anollx'i and iiKire extensive method of determining the per cent bagged has been possible through the correlation of the recorded grouse kill by counties'"' with county game habitat maps (which are completed for a number of counties in New York State). In addition to these two sources, it is also necessary to know the average density of the grouse population for the year in question. This is obtained friiin the field censuses on the various grouse survey areas. From the game habitat map is obtained the quantity of grouse cover. The components of this total are weighted according to quality to get the quantity of cover of a quality equiv- alent to that on the survev areas. To this is applied the grouse-per-acre figure from the field * CcDtUfet on Imth tlir Mirhican and Minnriuln sludirs wrre nnnir b>- King'B gridiron method. A Taken from the hunlrr't rrporti obuinrd upon iviuance of new licrnaes. AS A GROUSE HUNTER 375 survey results to get an estimate of the October grouse population in each county. The number of grouse killed as recorded by the hunter reports is then doubled to take care of such inaccuracies as are described on p. 372. The per cent bagged is then obtained by taking the ratio of grouse reported bagged to the estimated grouse population. These results are only a general indication of conditions and trends. The arbitrary assign- ment of cover quality values is obviously open to error, as well as other details of the system. But probably more significant than any of these is the possible error accruing to a faulty distribution of kill reports in relation to residence of the reporters. Thus, it is prob- able that the per cent bag for counties having large cities is exaggerated due to reports of many hunters who actually went outside the county for their hunting.* The take in seven counties from 1931 to 1934 ranged from 4 to 28 per cent. Half of the figures were between 10 and 20 per cent. The extreme range in a single year was 9 per cent. Considering the limitations of this method, the results corroborate the field studies remarkably well. Average Bag. Seasonal and Daily To those who are on the threshold of becoming grouse hunters, this topic will be a revela- tion. And to those "old timers" who may still be smarting from a bad day when they "couldn't get their eye on 'em", it should prove reassuring, for there are many more whose "luck" has been worse. There probably is no other American game bird which presents as difficult a target as does the ruffed grouse, except, of course, in those wilderness areas where the species has not yet become man-wary. That such a ganie.y bird should develop in the space of a century, is a tribute to its adaptability. The old journals are replete with records of large daily and seasonal takes, usually by market hunters. Forbush"" mentions several cases occurring in Massachusetts. He says. "I can remember when a market hunter going out from the city of Worcester by train each day, walking to the covers and returning at night, killed from 10 to 15 birds daily . . . Five men of Foxborough snared grouse prior to 1888 and averaged about TOO birds a week . . . Mr. George Howes shot and marketed 398 birds in one shooting season". Haight' ' relates of two men shooting 998 grouse from the 15th of September, to the following 31st of Jan- uary, a total of 122 hunting days or an average of four birds per man-day. Not only are the days of the market hunter gone but so also arc the days of such legal bags. Current efforts must necessarily be within bag limits of three a day and 15 a season in New York. An indication of the average seasonal bag of grouse per hunter in New York may be obtained from the reported kill of licensed sportsmen. These are given in table 5.S. Considering licensed grouse hunters as a whole, the maxinmm average seasonal bag dur- ing this period was approximately five grouse per hunter in the best year, and in the worst year, about one. Leopold records the average bag for all hunters in Wisconsin in 1931 to be .3 bird per hunter for the season. This is somewhat lower than the average for all New York hunters during 1923-1939. ,1';.'.'' Information on daily bag has been obtained on the field hunter-check studies in 1930 and 1931. In these two years the data show that it took 19 and 12.6 hours of hunting, respectively, per grouse bagged. The hunting "day" was 5.75 hours in 1930 and 3.4 hours in 1931. Reduced to an 8 hour day basis, the average daily bag per hunter was .42 grouse per day in 1930 and .63 in 1931. The actual "daily" bag in the short days which were * Reports are assumed to apply to county in which license was taken out. 376 INFLUENCE OF MAN recorded was .33 in 1930 and .37 in 1931. The bag of .63 grouse per 8 hour day in 1931 — based only on the efforts of grouse hunters, probably increased slightly with the increase of the birds in succeeding years. However, the check made on Conneiticul Hill in 19.36 only showed a slight increase to .7 grouse per 8 hour day and only .2 grouse for the average hunt- ing "day" of 2.1 hours. However, this low "day" is partly accounted for by the fact that only the part of the hunters' day that was spent hunting on the check area was counted. A measure of the distribution, rather than the size, of the daily bag may be obtained from the proportion of hunters taking one or more grouse per day. In the 1930 study this was 25.0 per cent; in 1931, 26.9 per cent; and in 1936. only 15.3 per cent. Thus, even restricting our take figures to hunters alone, in fact, to hunters in grouse cover, only a small proportion actually get grouse in any one hunting day. Crippung Loss The number of grouse crippled by hunters but not brought to bag, being an exceedingly difficult factor to evaluate, has always been the subject of wide differences of opinion. To some observers it has appeared negligible while to others it loomed even larger than the bag itself. The Investigation has had two opportunities to work on the problem, once in connec- tion with the hunter-check in 1930. 1931. and 1936, and again with the grouse population control ex])eriments in 1933-34, 1934-35 and 1935-36. An exaggerated version of the crippling loss is obtained by subtracting the hunter "take" from the difference between the pre-season and post-season censuses in a hunter-check sur- vey. This result represents the entire loss of birds during the hunting season except for those brought to bag and includes losses due to predation, disease, and accident as well as from hunter-crippling. It is assumed in these surveys that grouse movement into and away from the check areas is balanced. The results on the hunter-check areas indicate that the whole hunting season loss is approximately double the hunter take. These data are summarized in table 57. TABLE 57. ANALYSIS OF HUNTING SEASON LOSSES OF GROUSE IN NEW YORK— 1930, 1931 AND 1936 Tompkins county hunter ch^'clt areas PuUic hunting grounds 19.-)0 1931 1936 ISumbt-rof gruusc before the hunting st-ason Number of grouse after the hunting season. . Totiil loss of grouse during hunting season. . . Portion of hunting season h)ss of groustr 62 51 11 6 5 54.5 45.5 186 126 60 29 .31 48.3 51.7 94 69 25 13 I'ortion of hunting season loss of groutu* due to all other causes, including crippling by 12 Proportion of hunting season loss due to 52 0 Proportion of loaa due to all other causes. . . . 48.0 Thus, the losses due to all other causes during the hunting season approximately equals the loss due to hunter "take". Saunder's work in Michigan, which showed a hunter "take" of 6.7 per cent, gives a total Ocloher loss (which ])cri()d is over double the length of the open hunting season of 12 days) from all sources as 15.9 per cent of the September popula- AS A GROUSE HUNTER 377 tion. Covering as it does a period considerably in excess of the hunting season, the 9.1 per cent loss due to causes other than the hunter "take" is roughly comparable to the New York result. Trippensee's figures indicate a considerably different relation between "take" and cripples in Minnesota — his "other losses" amounting to only 21.6 per cent of the hunter "take" for the hunting period. How much of the 8.1 per cent and 16.7 per cent losses of pre-season populations from these several sources in New York can be attributed to crippling is undeterminable. The cen- suses on these areas were not sufficiently detailed to evaluate the effects of such factors as predation. These figures do represent, however, the upper limit of possible crippling on these areas. Beginning in the fall of 1933, certain portions of the Connecticut Hill survey area were set aside for grouse population-control experiments. This control consisted of collecting every bird possible on these portions the first year and in succeeding years collecting enough to reduce the grouse density to half the normal maximum. Coincident with the collecting of the birds, intensive census work was carried on to check on population changes due to birds moving into or out of the area. In so doing, it has been possible to get a reasonably accu- rate check on lost cripples by careful follow-up searches. While it would be perfectly possible to determine the percentage of the fall population that were collected, crippled-in-collecting, and died from predation. these figures would be mean- ingless in view of the length of the collecting period involved and the purpose of the work. The crippling therefore has been worked out only in relation to the "take". These data are summarized in table 58. These figures represent a minimum estimate of the numlier of birds crippled but not brought to bag. Included in them are the number of birds knocked down but lost and that subsequently may recover. It should also be remembered that the men making the collec- tions on which this table is based were relatively good grouse shots. Thus the figure repre- senting the number of grouse crippled but not bagged is probably lower than the average for 378 INFLUENCE OF MAN T.\BLE 58. CRIPPLING LOSS ON POPULATION CONTROL EXPERIMENT- CONNECTICUT HILL Year Number of grouse shot down Number of grouse bagged Number of grouse crippled but not bagged Per cent of shot birds that were crippled 1933-34 1934-35 1935-36 31 62 30 27 56 26 4 6 4 12.7 9.4 13.3 Total 123 109 14 11.4 the state. Considering the maximum number of birds that might have died following crippling (table 57) and the minimum estimate in the table that follows it, the Investigation believes that the probable loss from hunter crippling is normally about % of the birds shot down, or 2 per cent of the pre-season grouse population. Total Hunter Kill The sum of the hunter take and the loss from crippling gives us the total hunter kill. On the Tompkins County hunter-check areas in 1930 the number of birds that disappeared between September 1 and November 30 was 17.0 per cent while in 1931 it totalled 32.3 per cent. This included losses from all causes including hunting. A figure closer to the actual hunting loss is obtained by applying the estimated ratio of cripples as given in the preceding topic to the bag and adding that amount to the six and 29 birds bagged in 1930 and 1931 respectively. Then, in 1930, the estimated total kill was 11.3 per cent of the pre-season population and in 1931 (which is considered as a normal grouse hunting year for the increase phase of the grouse cycle) it was 17.7 per cent. The 1935 public hunting grounds hunter-check showed a total kill of 15.4 per cent. Since these mathematical figures are obviously only approxi- mations of conditions as a whole, we may summarily conclude that the total kill by hunters in New York is about 17 per cent of the prc-hunting season grouse population. Success Ratio Success ratio as herein used applies to the ratio of birds bagged to birds flushed by the individual hunters. The ratio may vary from 0 per rent to 100 per cent but the results iiliiaiiicd on the 1930 and 1931 hunter-checks as summarized in table 59 show that it is more iniiinioiilv closer to zero. T.VBLE 59. SUCCESS U.Vno l.N CRULSE UUNTlNG^TO.MlMvl.NS COUNTY .VNU PUBLIC HUNTING GROUND HUNTER- CHECK ARE\S NuuiIht of grouse Hushed Number of trrousf^ hnfigfd Year Snccejts ratio (per cent) 1930 68 :i<(7 ii; 6 29 i:t 8.8 1931 7.3 IM36 11.1 Thus the success ratio in 1930 was 8.8 per cent or one bird bagged to every 11.4 flushed and in 1931 was 7.3 |)cr cent or one bird bagged to every 13.7 flushed. Tlie 1936 check shows the highest ratio of all, 11.1 per cent or one bird bagged for every nine flushed. ^5 A GROUSE HUNTER 379 This ratio, being based on total birds flushed, is considerably smaller than the ratio of birds bagged to birds shot at which is the correct test of success in terms of shooting skill. This ratio was not determined on the 1930 and 1931 check areas but in 1936 it proved to be 27.1 per cent or about one bird bagged for every four shot at. Effect on Grouse Abundance of Man's Hunting Whether the hunters harvest 10 or 20 per cent of the grouse is of importance from the game management point of view, mainly as it affects the continuing abundance of the species. Under ideal conditions, the hunting loss should be the maximum that would leave an optimum breeding stock for the succeeding year. To illustrate, assuming an area with a good grouse population in the fall, there might be two birds of the year for every pair of breeders existing on the area the previous spring. In such a case, if there were no other sources of mortality from October through the next breeding season, the hunters could harvest two birds of every four, or 50 per cent, and still leave the optimum breeding stock. Actually, however, this figure represents the percentage from which must be deducted the anticipated winter loss due to other causes, in order to get the maximum allowable harvest by hunting. Examining the record in retrospect, the deductible figure has varied in different years from 11 to 60 per cent (the winter loss on the Connecticut Hill study area). Again assuming a fall population to which the above mentioned 50 ])er cent is aj)plicable, the theoretical permis- sible harvest would vary from 39 to 0 per cent of the pre-hunting season level. But pro- ductivity is not constant. On this area it has varied from year to year so that, instead of .50 per cent, the total allowable overwinter loss with respect to what may be considered an ade- quate breeding population has ranged from 17 per cent to 70 per cent. This variability is to a large extent compensated for. however, by the fact that high overwinter losses have tended to be associated with high fall densities and vice versa. Fortunately, too, there is substantial evidence which indicates that these figures may run much higher without danger*. The above "deductible figures" are based on an area where no hunting was allowed. Many of the birds that might have been taken by hunters were lost through other causes. If the) had been taken by hunters, it is improbable that in every case other birds would have been lost in their place. One may then fairly conclude that the 17 per cent hunting loss estimated does not endanger the continued abundance of the species in well-stocked coverts. That modern hunting is seldom an important factor in altering the natural course of grouse abundance is further indicated by the record in New York in recent years. Following two years of closed seasons during the low-abundance period of 1928 and 1929, the season was re-opened in 1930 when the abundance curve had recovered to about 30 per cent of its ultimate peak. Many were the disapproving cries of anxious sportsmen who, having become thoroughly alarmed by the "disappearance" of the birds in 1927, felt that it was not yet safe to allow an open season. But in spite of this and succeeding open seasons, the grouse continued to increase until a high level of abundance was reached, which, in general, has been maintained until the present (1942). To further indicate the part played by decimation from hunting, it is helpful to examine the relative importance of the various sources of loss by which the potential productivity of a spring grouse population is reduced. The charts included under the discussion of Net » See Chapter XII. p. 538. 380 INFLUENCE OF MAN Productivity* show graphically the fate befalling an average 1,000 grouse eggs and the cor- responding breeders under the major variations in survival pattern experienced during the Investigation. In each case "hunter take" accounts for but a small proportion of the total reduction of the potential 1,000 eggs. In figures 52, 53 and 54 losses prior to the ovenvinter period are the same and the 17 per cent attributed to hunting represents 38 birds (approximately 33 being bagged and five crippled but lost) comprising only about 4 per cent of the full year mortality. Wlien losses during the spring and summer are greater (figures 55 and 56) it becomes even smaller, both numerically and in proportion to the whole. In no instance depicted would elimination of shooting alter the population trend since no more than half of the birds so taken might be expected to survive if no hunting were done. Nevertheless it is obvious that as the season progresses the loss of the same number of individuals becomes of greater import. "'X% Wliili' occasionally an individual covert is adversely affected by overshooting, it must be concluded that llie general effect of hunting for sport as currently practiced on grouse, is not detrimental and may even have a positive beneficial effect during periods of grouse abun- dance by tending to reduce the opportunity for the agents causing periodic scarcities to become effective^. AS A HUNTER AND TRAIM'KK OF FREDATORSt The authors approach this subject with a good deal of trepidation. It is the object of the » Sm Chipter XH. p. 5.19. A It mutt be recognized that tlie discuision of this topic applief only to the iocrcjise aod pealc phases of K^ouse abundance. t See also Cbapler VII, p. 346. AS A HUNTER AND TRAPPER OF PREDATORS 381 Investigation to substantiate all conclusions with a foundation of observed or demonstrated facts, but man's relations to predators, by its very nature, must be strung together with num- erous postulations where facts and figures have not as yet been obtained. Our concern in this instance would be less marked were it not that the subject had already been treated at length by two divergent and highly-opinionated sections of the public. Statements made in progress reports of the Investigation in past years have in some cases occasioned outcries of varying intensities from both the "leftists" and the "rightists". Being damned by both extremes may possibly be interpreted as back-handed praise. Certainly it has instilled an even greater-than-normal feeling of caution. In all conclusions drawn, man's hunting of predators has been considered purely as it may influence grouse abundance. Number and Per Cent Taken by Hunting A reasonably accurate determination of either the numbers or per cent of predators taken by hunters is not easy. In the cases of some of the manmials, the license stub reports required of hunters and trappers in New York give valuable data although most of those reported are trapped rather than taken by hunting*. Only part of the predatory mammals, however, and none of the predatory birds are among the species which are reported to the Conserva- tion Department in this way. Thus, without this statistical help, the problem must be attacked in broad terms. Written records of sportsmen's club activities, bounty records (in those states which offer bounties), observations on hunter-habits and the relative vulnera- bility of various species to hunting by man must furnish the basis for discussion. In so far as their potential effect on grouse is concerned, the number of predatory species that need be considered is small. Among the mammals are the red and gray foxes, the New York and small brown weasels, and the skunk. Among the birds the great horned owl and Cooper's Hawk are the ones most important in New York, although the goshawk, sharp- shinned hawk and crow may at times become troublesome. The number of predators of any species taken by hunters depends not so much on the degree to which it destroys game as upon its vulnerability to man's weapons. That the crow is a much-hunted species and also of some importance as a grouse predator is largely acci- dental. That the horned owl is only occasionally taken by hunters, even though it is an'impor- tant grouse predator, is mainly due to its seclusive and nocturnal habits. That the red squir- rel is seldom important as a grouse predator and yet is conimonlv shot by hunters is due to the ease with which they are sought out and killed. With notable exceptions, man's hunting of predators has not resulted in their great reduction. Rather, it has forced the various species to become more wary and adept at avoiding man. In some cases, largely due to man's improvement of the environment for them, species have actually increased from former num- * See discussion of Man As a Tr-ipper of Fur Species, p. 382. 382 INFLUENCE OF MAN bers in the face of constant hunting. One must conclude that man. as a hunter of predators, is of little consequence to grouse conservation. Whether it be the normal, haphazard, "'catch-as-catch-can" shooting, organized "vermin hunts" or hunting for bounty, the numbers of important grouse predators taken by shooting are small. As for assisting in the management of grouse , sportsmen's efforts as predator hunters are relatively ineffectual. As A Trapper of Fur Species Trapping for fur is an economic enterpri-ie. Returns come in payment for the pelts taken and, in some cases, additional revenue from bounties and from carcasses. Several of the valuable fur-bearing animals are also important grouse predators. Because of the dual role of these species, trappers unintentionally (in contrast to predator hunters) exert an indi- rect influence on grouse. Fur bearers in this State which are also important grouse predators are niainh the red fox, gray fox, small brown and New York weasels and skunk. The raccoon, while a less im- portant grouse predator, is also trapped in some numbers. The take of all these species, ex- cept the weasels, has been recorded in New York since 1918, along with the reports required of game taken. Table 60 summarizes these data beginning in 1926. TABLE 60. REPORTED TAKE OF CERTAIN PREDATORY FUR BEARERS IN NEW YORK— 1926-1939* Number of Animals Reported Taken by Trappers and Hunters Year Red fox Gray fox Skunk Raccoon 1926 1927 18.726 35.658 10.616 10.913 8.644 8.118 7.697 9,278 9.009 11.229 10.593 15.550 19.452 25.852 2.402 2.338 2.269 2.775 2.652 2.806 3.443 3,527 4,933 4 757 4,247 4.9K0 5,250 6.975 73,648 120,002 72,623 74,850 53,541 56.338 44.905 51.063 32.988 33.247 42.669 36,701 33,952 41,170 28,750 40,908 1928 27,886 1929 24.23() 1930 20.820 1931 18.813 1932 18.686 1933 13.939 1934 20.364 1935 23.427 1936 26.921 1937 24,190 1938. 25,345 1939 27,459 Average 14.381 3.811 54,121 24.410 * The recorded take of otlier predatory fur-bearers not appreciably affecting grouse is not included here. Considered from the standpoint of the predators themselves, these data bring out several significant points. With red fox, skunk and raccoon, the take depends mainly on fur prices. It was very high during the booming fur market of the middle *20s. Then it drojjped mark- edly with the economic depression of the early '30s following which it has commenced to increase again. No species has shown any indication of a marked decline in number, or. as a corollary, pelt trapping of these species seems to take only the surplus population, if that much. It is apparent that ordinary fur tra|)|)ing h.is not resulted in any widesjiread reduction of any of these species. .An exception miglit occur during a ])roli>Mgc(l period of high fur prices, a circumstance not likely to occur. A substitute for high fur |)rices is the granting of a bounty in addition to the return from the |)ell. Pennsylvania has attempted control by this means but it likewise has failed to control any species except the bobcat. AS A F4RMER 383 Gersteir^ concluded : "As a predator control measure, the payment of bounties has proven generally inefficient as it has placed under control only one relatively small species popula- tion, while its effect on five others has been negligible". We thus conclude that even with the stimulation of adequate bounties, control of the important ruffed grouse predators is not accomplished in the sense of marked reduction in populations. Even if populations of fur-bearing predators are not reduced by commercial trapping or trapping for bounty, will not the numbers that actually are taken benefit the ruffed grouse population? In some cases they may. The taking of fox and weasel for fur may lower grouse nest mortality in localities where those predators are severely reduced and thereby increase the fall surplus*. This sort of trapping, however, is not commonly associated with the better grouse coverts of the state. Furthermore, it is a dubious benefit unless the added surplus is taken by hunting. Gerstell, speaking of Pennsylvania's experience with the bounty, concludes, "It has been impossible to prove that the operation of the bounty system over a relatively long period of years has improved game conditions. Furthermore, it was shown that the annual amount of money expended for bounty payments was controlled not by the abundance of predators, but principally by climatic and general economic conditions". The trapping of predatory species for fur, with or without the added economic stimulus of a bounty, is negligible in its effect on grouse abundance. AS A FARMER As one delves deeper into the complexities of the grouse problem, he becomes ever more impressed with the role that man plays in affecting the birds' habitat, .\ever is this more significant than in his work as a farmer. In the first place, at least in the Northeast, the most productive coverts occur in those portions of the range where the woodlands are broken up by considerable areas of land in farms. The farmer is responsible for this condi- tion. Secondly, since these lands are privately owned, whatever is done that affects grouse either for good or bad will be done by the farmer. He clears the land of woods, or keeps it clear. He controls livestock that may impair or destroy cover values. When he turns lumberman and takes products from the farm wood- land, he is again functioning as a manager of the cover that may or mav not support grouse, depending upon how he does his work. Clearing of Land One of the most generally accepted legends of primeval conditions in the northeastern states is the great abundance of game in the unbroken wilderness. Originating in the tales of the early settlers who found adequate supplies of game around their newly-made clearings, it was naturally assumed that this abundance was an attribute of the whole wilderness. Little did these pioneers realize that their very efforts in creating the clearings was largely respon- sible for the local abundance of grouse, deer, and turkeys which furnished them with a ready supply of fresh meat. Thus the clearing of the timber to make way for farming was the first game management technique practiced, however unwittingly, by the white man in America. But, as with all good things, too much is no good. The clearing of the land in many regions soon passed the optimum condition for grouse and, in pla'-e of the improvement of the coverts, the needs of farming caused a rapid destruction of the coverts which delimited the available * See Chapter VH, p. 346. 384 INFLUENCE OF MAN grouse range. In those flat, fertile areas of New York which proved to be excellentl)' adapted to farming, the grouse was exterminated and remains so today except for occasional "islands" of wooded swamp or otherwise untillable land. In the southern-tier counties, which are typical of the better grouse range in New York, the timber on the hill land was generally cleared on from 40 to 60 per cent of the area. It is here that one today can measure the effect of land clearing on grouse. Even when one includes all of the open land in the grouse range, the population density far exceeds that on wilderness areas of unbroken coverts such as the Adirondack mountains. Thus on the Connecticut Hill study area where the brush and woodland cover occupied about half of the land surface, the grouse density for the range (including all open land) has been from 22 per cent to 88 per cent above that on the Adirondack study area (which is 92 per cent wood- land and brush) and has averaged 50 per cent higher during the period of the Investigation. We must thus credit, at least partially, the farmers' land clearing operations with enabling an increase of one-half in the grouse productivity of the land on this type of range — and of producing that crop on only half the formerly available range. How close this condition lies to the optimum effect of land clearing is not known but it seems probable that a much higher productivity could be secured with from 10 per cent to 25 per cent of land cleared — provided it was well scattered in small units through the range. The arrangement of the open land on the range is more important than the quantity. Maintenance of Openings As just noted, the original clearing of portions of the land exerted a profound effect upon its utility as grouse range. The continued farming of tliese cleared lands has been equally important in maintaining the newly created carrying capacities of the range, some improved and some completely destroyed for grouse. On the intensely farmed lands, the effect has been to prevent any potential encroachment of the grouse on the destroyed range. On the half-cleared lands, this maintenance of the openings has made of the farmer a game manager. In recent years, the trend toward abandomnent of the lands which have proven submar- ginal to agriculture, has caused marked improvement of the grouse range. The release from tillage of these fringes of open land has created brushy areas which add valuable food- producing coverts to the range, increasing both the quantity and quality of the grouse cov- erts. If this abandonment is not carried too far, if a suflicicnt quantity of open land is main- tained well-scattered over the range, this improvement will be maintained. But in so far as the abandonment of open land maintenance becomes extensive, that range is destined, in the future, to aiii)roach the relatively low grouse productivity of the original wilderness. Only government purchase and management can remedy this condition unless other uses profit- able to private enterprise are developed. Pasturing of Livestock As a general proposition, the pasturing of farm livestock in New York is of little ini|)or- tance to grouse. It is fortunate that even where dairying is a major farm industry, the farmers do not practice woodland pasturing extensi\ely in the better grouse range, for rarely do we find woodlands heavily pastured even when cattle have access to them. Woodlots in the intensively farmed areas are conunonly [>a.-lure(l to the point of creating a visible "cattle line". Any grouse range so pastured is rendered almost wholly unproduc- tive of grouse regardless of its other attributes. Fortunately, this occurs mainly on small AS A LUMBERMAN 385 woodlots in areas which are unimportant as grouse range anyway. The Investigation has not made censuses on comparable pastured and unpastured grouse range and hence can not present specific figures on the effect of livestock pasturing on pro- ductivity or carrying capacity. The conclusions reached are based upon general observa- tions, which, however, have been very clear. It should be noted, in passing, that livestock pasturing, as well as the plow, is a tool of the farmer for maintaining open land. As such, it often assists in maintaining a high grouse population in adjacent coverts made possible by that open land. Livestock occasionally destroy grouse nests by trampling, though the loss from such a cause is not of great importance. Other Domestic Stock While the pasturing of livestock is the most important relationship of farm animals to grouse, chickens and other domestic fowl, dogs and cats in some instances may have some significance. A grouse nest was examined in Delaware Count) in which a chicken had laid eggs and caused the grouse to desert. This is not a common occurrence but worthy of passing note. In the summer of 193G, a grouse bmod on the Connecticut Hill study area was flushed from the midst of a group of chickens, all of them having been scratching about in the leaves together. Such contacts of grouse with chickens or other domestic fowl are not uncommon and may provide an occasional means of transmission of disease*. It is clear, too, that even without contact, traversing the same terrain as domestic fowl may possibly lead to disease. Dogs and cats, especially those allowed to roam at will, often become serious predators. The dog exerts the most effect in nest-destruction while the cat is adept at catching young birds and occasionally adults. The im|)ortance of these domestic animals as grouse predators is detailed in Chapter VII. AS A LUMBERMAN'^' When the white man arrived, he found what is now New York covered with a vast forest, broken only by water areas, marshes, cliffs, occasional blow-downs and the scattered clear- ings of the Indians. Forest cover types merged into one another with a minimum of definite edges, except where such natural forces as wind, fire, insects, disease or decadence had broken the woodland canopy. Undisturbed by civilized man, the forest had achieved a stage * See Chapter X, p. 415. A By David B. Cook. 386 INFLUENCE OF MAN of development approaching the climax. Areas occupied by the earlier successional stages so necessary to high grouse production were probably few, small, and scattered. One of the first products taken from the forest by the white men were heavy planks, sawed from the choicest white pines. Later they cut oak and other hardwoods, both for their own use and for export. Because of the superabundant supply, these early cuttings were light and highly selective, taking only the largest and choicest trees of certain species. Probably, such operations disturbed the forest very little. Few openings big enough to seriously affect the forest composition were made. As population increased and transportation facilities improved, the forests were exploited for a greater quantity and a wider variety of products. I\ot only choice saw-timber but lower grades of logs and wood were extracted to meet the growing demand. In the Adirondacks, where river driving was the chief means of transport, only softwoods were cut. In the south- ern tier, tremendous quantities of pine were sawed up and the lumber rafted to market. Untold millions of hemlock trees were stripped of their bark for tanning and left to rot in the woods. In all but the oldest-inhabited parts of the State, the lumberman and his sawmill kept just ahead of the farmer and his grist mill. Generally, these cuttings were heavy enough to break up llie forest canopy. The removal of the bigger trees, with the attendant increase of sunlight and air, the ripping up of the soil in skidding and roadbuilding, and the effects of occa- sional fires set back the course of forest succession and brought in many light-demanding trees and shrubs which produce grouse food. Thus edges were produced in abundance as each year's cut followed the last. Later, with decreasing limits of merchantability, the tendency was to clear-cut extensive areas. This was especially true in the Adirondacks, where large bodies of spruce and fir were cut for pulpwood; in the southwestern Catskills, where hardwoods were clear-cut on a short rotation for acid-wood; and in the western part of the southern tier, where timber- lands were stripped for mine props. Elsewhere, the portable sawmill operators had taken off even the smaller trees. Such lumbering operations tended to break up the pattern of the forest into a multitude of small blocks of varying age. They stimulated the growth of advance reproduction, under- growth and herbs. Especially where fire intervened, the climax forest composition was replaced by woods representing earlier stages in the successional series. Abundant grouse food-producing plants are usually to be found in these. Thus, the vast and uniform areas of virgin forest were replaced by a patchwork of small lots of varying age. composition and density. Such a forest paltcrn, with its varied and productive flora and its tremendous amount of edges, is ideal for grouse. The now-familiar cover requirements — s|)ring breeding grounds, summer and fall feeding grounds and winter shelter — were available everywhere. As fast as new areas of suitable habitat developed, the grouse moved in and occupied it. Though per- haps losing a little territory through fire or too-close cutting, they gained immeasurably in numbers, as is attested by early records of abundance. With the disappearance of "inexhaustible" supplies of virgin tinilicr, luinhering is now carried on in smaller units. Trees of less size are also in demand. The production of such second-growth, though still largeh accidental, does rccpiire some care on the part of the land- owner. There are taxes to be paid, fires to be kept out. As the value of land and timber AS A CONSERVATIONIST 387 has increased, so, too, has the intensity of forest management. Silviculture, the deliberate cul- tivation of the forest, is gradually replacing the old, haphazard methods of exploitation. Such necessary features of lumbering as roads, skid-trails, rollways and piling-grounds are very useful to grouse. These grow up to tangles of herbs and shrubs, especially briars, which persist despite occasional trampling. Thus the axe, like the plow, has exerted at times a favorable and again an unfavorable influence on the habitat, on the condition of which, in the long run, the number of grouse largely depends. AS A CONSERVATIONIST In our system of free private enterprise it has taken society a long time to recognize the need for conserving our vital resources. True, from colonial times there have been a few far-sighted individuals who have warned of the consequences of continued exploitation, but their voices were crying in the wilderness. In more recent times these consequences have become all too apparent and for many the time for retribution has arrived. The decline of millions of acres of once-productive land, and with it much of its former crop of wildlife, has resulted in the abandonment of vast areas of farm and forest. The concept of conservation of renewable resources — meaning %vise use — as a practicing credo of man has gained much acceptance in recent years. This trend toward a more per- manent system of land management has not been sudden, although its progress has acceler- ated since the turn of the century. Since the first law restricting the unlimited pursuit of grouse, there has been a gradual evolution of effort toward its conservation. Man himself has evolved as a conservationist. First believing that the enactment of laws restricting the take of grouse would adequately maintain a satisfactory grouse crop, he has gradually broadened the scope of conservation effort. Today the emphasis revolves around 388 INFLVEI^CE OF MA\ tlie practices of soiiiul laiir] management. The passage of laws restricting the activity of hunters was not only the first effort at con- servation of grouse, but also was a forerunner in the field of cooperative effort in conserva- tion. Restocking of coverts, establishment of refuges and bounty pavments for destroying predators are later developments of cooperative conservation practice. The basic practices of good land management, however, remain largely a matter of individual concern and their use or their disregard, an indi\idual determination. The ()i)portunity for the further development of man as a conservationist, especially in the collaborative sense, is great. Why should there not be community cooperation in protect- ing coverts from fire and grazing; in making plantations for new cover; in the marketing of woods products that will permit more productive woodland management methods for both trees and grouse? Therein lies future progress. Laws In the early years of the white man's colonization of the northeastern states, the ratio of hunters to grouse was so low that the supply seemed inexhaustible. No particular cognizance was taken except the harvesting of them for food. There was no need for protection and no conception of a possible need for cultivation. But as the human population increased and commercial facilities became available, lean years of grouse harvest appeared, likely being years following large losses. Understanding only the loss due to hunting, man turned naturally to a means of checking that loss by controlling hunting, by passing laws restricting the liber- ties of the individual. In New York the first law on grouse was passed in 1791. There followed a gradual spread and increase in hunting restrictions. These included the elimination of hunting during cer- tain seasons of the year, the curb on and elimination of market hunting, the prevention of sale, limitations . 630. AS A CONSERVATIONIST 391 New York today the less destructive avian predators are now protected by law. The mam- malian predators, with the exception of the foxes, weasels, red squirrels and house cats, are protected except during the prime fur season. Many of these restrictions, especially those protecting various species of hawks and owls, are largely unenforceable. They depend primarily for effectiveness on cooperation from hunters, which in turn depends upon their "education" for its degree of success. Any attempt to estimate the indirect effect these laws have on grouse is mere conjecture. It is the opinion of the Investigation that it is small, whether the law protects the predator or not. Grouse mortality in the Northeast is to a considerable extent the immediate result of predation but most of this loss seems destined to occur anyway. The primary values accruing from protection of predators are other than saving game. Refuges, Sanctuaries and Land Posting A study of the value of a refuge as a game producing unit in comparison with a public hunting area was made on the Pharsalia Game Refuge and Chenango Public Shooting Grounds Area in central New York. The refuge had been established for a period of five years prior to the beginning of the study. Public shooting grounds had been open to public shooting as private land prior to the study and had been marked as "Public Hunting Grounds" for a year when the study began in the winter of 1935. Hunters commonly frequent this region from the nearby cities of Binghamfon. Cortland and Norwich as well as from numerous small communities in the vicinity. \^Tiile no field check on the hunting pressure was made, the area lies in one of the best upland hunting sec- tions of the state. The hunting pressure was at least average on this area. The public hunt- ing ground was but 3 per cent larger than the refuge, thus the data obtained was essentially comparable from the quantitative standpoint. A comparison of the amount and distribution of the types indicated no apparent impor- tant differences. The cover within each area was essentially independent of surrounding lands. Thus, the two survey areas may be considered for all practical purposes comparable subject only to a 3 per cent correction for size. Three censuses were made on these areas, all in successive winters, the first being in 193.S. The data on the grouse populations for the refuge and check area are summarized in table 61. TABLK 61. UUFFED GROUSE D\TA— PHARSALIA REFUGE AND CHENANGO PUBLIC HUNTING GROUND SURVEYS— FEBRUARY 1935-1937 Year OinTtilioiis 1935 1936 1937 Rpfuge Area Check Area Refuge .Area Check Area Refuge Area Check Area 438 740 270 11 166 8.1 9.0 537 765 298 9 252 5.5 6.0 301 257 231 24 134 10.0 11.1 289 592 723 21 1.35 10.2 11.2 341 480 247 117 11.5 12.7 465 489 338 5 163 8.4 Acres of coverts per grouse 9.3 392 INFLUENCE OF MAN In 1935. the Refuge had only 66 per cent as many grouse as the public shooting area while in 1936 the populations were practically identical. Then in 1937, while the grouse population on the former continued to decrease, that on the latter increased until it carried 39 per cent more birds than did the Refuge. While the densities of numbers have essentially the same ratios as the populations, the 1936 population densities on both areas were considerably lower than those of the previous year. But in 1937 the density of grouse on the Public Shooting Ground showed a marked increase while that on the Refuge continued to decline. The analysis of the effect of the small differences in the areas may serve to explain some of the variations in grouse num- bers, but the vital fact still remains that the protection afforded by the Refuge during the period concerned did not serve to enlarge the crop of grouse. The conclusion seems inevitable that refuges are of little value in maintaining popula- tions of ruffed grouse when hunting pressure is not excessive. Fluctuations continue regard- less of the protected or unprotected status of the coverts. A sanctuary is merely an undeveloped refuge. Its value as a means of producing grouse, if different from a refuge, is generally less. Yet' we must not overlook the possible value of a refuge or sanctuary for preserving seed stock on areas that have an abnormallv high hunt- ing pressure, as on those near large cities. Rut such areas only function to prevent extermi- nation and not as a means of furnishing hunting on surrounding lands or of increasing the grouse population on the areas themselves. The practice of land posting by private land owners, if done objectively to improve the grouse conditions, is gcnerallv unwarranted. Its effect is the same as that of a refuge or sanctuary. de|)ending upon whether the area is utilized and managed or held inviolate. If the object of posting is to keep the hunting privilege for the owner or to prevent trespass, the problem is outside the province of this discussion. The explanation of the relative lack of usefulness of the refuge to grouse appears to be three-fold. Violently cyclic species can not exceed definite population levels, nor can they long maintain the maximum levels they do reach. The degree of sedentariness of a species tends to be in inverse ratio to its adaptability to the refuge principle, and grouse are quite sedentary. The adaptability of a species to management by the refuge principle depends upon the ease with which the species adjusts itself to recognize protected areas. The ruffed grouse is not very adaptable in this respect, as contrasted with the ducks, for example. Added together, the facts indicate that llie refuge may serve to prevent extermination of some grouse in areas that might be overshot but that it is seldom a priinarv tool of the grouse manager for increasing the crop. EFFECT OF MAN'S AIDS ON GROUSE The tools of the white man's civilization have exerted a most profound induencc upon the grouse since the first colonization of the Northeast. Ironical as it seems, most of these aids have worked both for the benefit and detriment of the species, depending upon how and to what extent used. While man in his various capacities has already been discussed, it seems j)ertincnt at this point to consider further the effects of his implements on grouse. Consid- ered somewhat in their proper chronological sequence, the more important ones are the axe and plow, types of guns, traps and snares, fire, domestic animals, automobiles and high- ways. EFFECT OF MAN'S AIDS ON GROUSE 393 The Axe and Plow If remaining areas of virgin wilderness may be accepted as comparable with primeval conditions, grouse populations in pre-colonization times attained but relatively sparse densi- ties. Theoretical consideration of the probable status of early grouse populations leads to the same conclusion. The grouse is a species largely dependent upon cover type edges for the attainment of large numbers. Edges must have been scarce in the climax forest, existing only at the border of different climax types, along water courses, bordering burns set by lightning and around Indian clearings. Hence, the carrying capacity of the original range was low compared with the species saturation level. Grouse being an "edge" species, the clearing of the land with axe and plow increased the carrying capacity of the range. Woodland borders were created mile upon mile as the embryo empire pushed back the wilderness border. Coverts were broken up and diversified. Except where vast areas of fertile land induced continuous land clearing, the effect was advantageous to the grouse. The plow followed the axe and to a considerable extent made permanent its changes. As a means of maintaining these new margins, the plow has continued to play an important role in preventing the return of the contituious forest. In many areas, the optimum balance for grouse of open and wooded land was jiasscd and the axe and plow then jiroduccd an immense reduction of available grouse range. Thus being the tools which cleared off the forest and maintained the open land, they have been of first importance both in improving grouse range and in destroying it. As time went on, the bread basket of the nation shifted westward and much of the land which had been cleared for crops in the Northeast proved unprofitable for farming. This decline of agriculture on subniarginal lands has led to considerable abandi>nmciit. Uidess carried so far as to obliterate most open or semi-open spaces, this trend is a distinct advan- tage to grouse. The axe must, in the future, be one of the chief tools through which the wildlife manager will hold and improve these grouse lands. The plow will also j)lav its j)arl in maintaining the edges so necessary to good grouse range. Fire Records show that most fires occurring in woodland are man-set. either intentionallv or accidentally. Thus the effect of fires upon the grouse is largely the effect of one of man's tools. With the efficient control now operating throughout most of the Northeast, forest fires are generally confined to small areas. They have little effect on the birds directly, although, when occurring in the spring, they may cause appreciable losses to nests. It seems rather ironic, but small woodland burns may actually improve grouse cover by stimulating growths of food-bearing trees and shrubs. Phillips'" states that the highest concentration of grouse he ever observed was in an area which had been burned over a few years before. While small occasional fires may be beneficial to grouse, extensive and repeated burning results in abandonment of the habitat for many years. In portions of the grouse's former range, notably in the Ozarks, annual burning for the "improvement" of the range for live- stock, plus overgrazing, has resulted in the extermination of the species. 394 INFLUENCE OF MAN Automobiles and Highways Before the advent of the motor car, grouse hunting was largely confined to an area of a few square miles near home which could be reached on foot, or a few selected coverts not many miles away that were accessible by a short rail trip or a buggy ride. Today, a day's hunting may involve two hundred or more miles of motoring and hasty inspection of a num- ber of coverts where the sportsman knows, from his own experience or his guide's, that there are some grouse and that he is likely to find them unless they have been disturbed or taken by other hunters. Modern highways and automobiles have opened distant coverts to the citv dweller. Coincident with the development of these arterial routes in the Northeast, many of the old dirt roads, which proved to be the "back roads" of today, have become abandoned. Areas once served by these roads have become inaccessible to modern cars. So, two changes have been working in our road systems which tend to counterbalance each other. The hunter's mobility has been increased extensively but has decreased locally. The net effect is difficult to evaluate but, in itself, has probablv had little influence on grouse num- bers. Whereas, formerly, hunters stayed "in their own back yards," they now are in someone else's back yard. Except that the total number of hunters has greatlv increased, the back yards are used about as formerly. Roads and their accompanying poles and wires, affect the grouse in other ways than afford- ing transportation for man. Collision with wires results in some mortalitv. particularly in the fall "crazy flight" period, and a few birds are killed by automobiles. Roads are openings and, when running adjacent to grouse habitat, may encourage concentrations at the edges. This tendency is enhanced when the road ditches and margins are allowed to grow up to low shrubs and vines. Added to this, the road itself, if of dirt or gravel, provides a source of grit and a place for dust bathing. Whether this is a desirable condition or not depends upon how concentrated the birds have become and upon whether or not hunters take too great an advantage of such concentrations and overshoot the area. Generally, this is unlikelv to hap- pen and roads as openings actually serve a very useful purpose. Guns, Traps and Snares llic old-fashioned sportsman paid more attention to the bobwhitc, woodcock and heath hen — birds that lay better to his well trained dogs — than to that erratic bombshell, the part- ridge. In those days, the nineteenth century in particular, it was the market hunter rather than the sportsman who reaped the largest harvest of grouse. Throughout most of the North- east, market hunting flourished until late in the century. With few restrictions the birds were taken with guns, traps or snares and brought to market in the cities. While the guns of this era were not as efficient as are (he nioie modern weapons, thev were nevertheless very effective on the relatively "uneducated" grouse of those days. For the same reason, traps and snares, while less effective on the wary birds of today, were also very efficient. This market hunting was business, not spoil, and ihe tools utilized had to be effi- lieiit if the business were to be successful. Improvements in weapons have been at least |)aili;illy ollset by adaptations of the liird itself to this change. Phillips'™ states: "Technical im|)r()vements in the past forty years we can set aside, for the early hammer guns, in skillful hands, were nearly as effective as the more dainty modern weapons." The inference will lie gathered that possibly the shooter himself EFFECT OF MAN'S AIDS ON GROUSE 395 has changed. He continues, "if we compare the average skill of the present day brush shooter with the average of the market shooter fifty years ago, the result would be highly amusing and very disastrous to the pride of the modern edition". Phillips then points out that the great increase in number of hunters in recent years makes the aggregate "as potentially destruc- tive as he of the old school". Increased protection for the birds and improving coverts combined with the response of the grouse to more intensive hunting by gaining steadily in his ability to steer clear of shot, have balanced the growing popularity of the sport with its improved weapons so that serious depletion from hunting is not likely to occur. Grouse Dogs* There has been an historical sequence in the employment of dogs as grouse hunting aides which must be laid alongside the accounts of other factors affecting grouse to make this report complete. The authors have also had an opportunity to attempt the use of dogs as an aid to game surveying, which has made possible some observations of general interest. Further- more, to many sportsmen, grouse dogs are inseparably associated with grouse hunting, the perpetuation of which was a prime motive for this Investigation. Hence this special sec- tion on grouse dogs. * By J. Virlor Skift. 396 INFLUENCE OF MAN In the early days, probably the first dogs to be used in partridge hunting were of the spaniel type. These were trained to flush grouse into a tree and '"bark them up" until the hunter could arrive and shoot them down, one by one, starting with the lowest to avoid flush- ing the rest. Alexander Wilson'"", 1812, cites the advantage of a "good dog": "the more noise he keeps up, seems the more to confuse and stupify them, so that they may be shot down, one by one, till the whole are killed." In some of the remaining wilderness areas of Maine and Canada, this type of dog and hunting are still occasionally employed. As grouse characteristics began to change in adaptation to man's influence, this style of hunting became impractical — grouse became too wary of both man and dog. Large forested areas, where it could best be done, gave way, throughout most of New England, to discon- nected coverts of small woodlots, brushlots, orchards and hedgerows. Thereupon developed a golden era in die history of grouse hunting over pointing dogs with the setters, notably the Gordon, emerging as the undisputed favorites for many years. If every dog does have its day — this was surely the setter's. Strains of grouse hunting setters were zealously guarded through careful breeding and selection and passed on from father to son. Only those which had a great nose, and could be trained to hunt close-in, were considered. It is probable that even then the patient, painstaking training of a good grouse dog surpassed the education given our finest field trial champions of today. Credit for developing these great dogs must be divided among the "sports" who hunted for fun, and the market hunters who had to have a dog with which they could deliver the goods day in and day out. It was this era which produced so many of the great grouse dogs, grouse hunters, grouse hunts and grouse hunting stories, and it was a sad day when old timers began to realize that it was coming to a close, thirty or forty years ago. Various reasons have been cited but four are given most frequently — increased wariness in the bird with accompanying reluctance to "lay to the dog"; the introduction of the pheasant which spoiled many good grouse dogs and grouse hunters; less breeding for the field and more for the bench and trials which resulted in changed canine ideals which were not appreciated in the thick grouse coverts; and periodic declines in grouse which were so severe as to discourage the breeding and owner- ship of specialized grouse dogs. Probably all four operated in varying degrees in various sec- tions of the Northeast. Towards tlie end of this period the English pointer began to take its place in the sun as a grouse hunter. Its relative merits were argued at great length in sporting publications of that day, even as thov still are today. It had one distinct advantai;c in that it was shorthaired and did not accunmlate burrs hke the shaggy setter. Generally, it was also less timid and less inclined to gun-shyness. The decline in specialized grouse dogs of the pointing type continued over two or three decades and up until very recent years. The number of grouse hunters actually increased but a greater prdjxirtiiin hunted without dogs or turned again to all-around types such as the cocker or springer spaniel. These types could be made to hunt close and were particularly useful in finding dead or wounded birds. They were also more adaptable to hunting for a mixed bag of pheasants, grouse, woodcock or even rabbits and s(]uirr('ls. It has come to be a well recognized fact that few pointing dogs, except in llicir "Id age, can be successfully switched from pheasant hunting to grouse hunting, or vice-versa. It is too much to ask the average dog to range wide and fast over open country for pheasants one day and then to slick to the cover and hunt close for partridge on the next. In very recent years, it is possible that we may have entered what will prove to be another EFFECT OF MAN'S AIDS ON GROUSE 397 era in grouse-dog history. Skeet and similar clay bird shooting have produced many splendid wing shots who are turning to partridge as the top gunning bird of the northeast. This, in turn, has revived interest in grouse dogs of all kinds. Along with this development is the happy circumstance that agriculture has long since passed its peak in the northeast and each year finds thousands of acres of cleared land reverting to brush and — grouse cover. History will record the outcome of these two highly significant trends. To the old line spaniels, setters, and pointers has been added a newcomer — the highly adaptable, all purpose, German shorthaired pointer. Although not fully tested, it gives promise of great things as a cover dog. It is especially tractable, smart and its extra dash of hound blood gives it a great instinct for hunting. However, it is not as stylish as the setter or pointer and its usual dark color is a great handicap in close cover where it becomes vir- tually impossible to hunt without a bell. Its chief advantage seems to be that it is more easily broken and handled by the amateur. Other largely untried breeds may also merit explora- tion. Among experienced grouse dog men, those who are able to lay aside strong personal likes and dislikes as to breeds or strains generally agree that, today, the making of a good grouse dog is more a matter of the individual dog (what he has in him inherently) and the training he receives, than it is a matter of a certain breed or strain. However, breeds and strains do have some generally well recognized characteristics and it behooves the uninitiated prospec- tive buyer of a grouse dog to ask himself just what it is he wants his dog to be able to do and to keep in mind just where and under what conditions he expects him to do it. A thought- ful consideration of these items will help to indicate the best type or breed for a given job. As to the broad question — dog or no dog — ^the authors leave that one to the hot stove leagues and publications of the sporting world. The authors themselves are divided. If pressed for an answer it would have to be — a good dog, yes; anything less than that, no. The writers have naturally known many grouse hunters and seen many alleged grouse dogs in action. All too few of the latter have been properly bred, trained and handled. The answer to this question also hinges on the type of country to be hunted. In places where a sparse grouse population is under great hunting pressure, a dog is of doubtful value, except perhaps as a retriever. In such situations grouse become so wary that dogs are of little value as pointers or finders. They are also of doubtful value in the big woods wil- derness areas. In thick brushy coverts where birds are reasonably plentiful and the hunting pressure moderate, their value is almost unquestioned. In the early days of the Investigation, the authors attempted the use of grouse dogs as an aid to its field work on areas where birds were being followed closely throughout every season of the year. Tests were made on the Connecticut Hill area with the best grouse dogs locally available. Each dog was handled by his own master. Somewhat to the surprise of their masters, and to the disappointment of all, even the best of the dogs failed to do any kind of a job under spring, summer and winter conditions. Even in the fall, the fact that dogs have "good" days and "bad" days, made results anything but comparable. In the spring, scenting conditions are so exceptionally poor that it is not unusual to have a good dog pass within a very few feet of a nesting grouse and never detect its presence. In the summer, conditions of heat and rank vegetation quickly exhaust the most stout-hearted and best conditioned dogs, rendering them virtually inoperative. In the winter, increased tree roosting and snow roosting take the birds out of the scenting range of dogs; and very deep snow or soft crust, shackle a dog very effectively. 398 INFLUENCE OF MAN However, even though dogs cannot be relied upon as a tool for painstaking field work, the authors are, nevertheless, firm in their belief that the individual hunter can learn a great deal about the habits of grouse during the fall season by the close observation of a good grouse dog. Man, as a grouse hunter, is handicapped because he can' only see and hear and does not have a nose tuned for bird scent. Thus, most hunters who "walk them up" can only learn by observing and, to a lesser extent, by hearing. This reveals little about the bird's move- ments up to the point it explodes from the ground or a tree. A keen hunter, especially if he raises and trains his own pup, can learn more from his dog than he can teach it. One of the first and most important lessons has to do with that critical period between the time a bird senses your approach and the moment he actually flushes. It is no accident that a grouse usually flushes from the other side of a piece of brush, a tree, a stone wall or whatever kind of cover is available. He has maneuvered himself into that position. A pup will often come to a staunch point and hold it expectantly only to have a partridge get up a long ways ahead and, usually, beyond a screen of cover. The pup and his master soon learn that she was pointing where a bird had been when first approached. In the meantime the bird had quietly sneaked ahead until able to fly safely away. As a dog gains in expe- rience, the first point is often momentary, followed by a series of starts and stops along a partridge's trail until the final point is made, indicating that the bird has reached his jumping off place. By fixing one's eyes on a point well ahead of the dog it is often possible to guess where that jumping off place is and then for the hunter to approach from a converging angle in such a way as to disorganize the bird's strategy and force him to jump so that he will offer a fair shot. It is usually a mistake, if the cover is at all heavy, to walk directly ahead of a dog on a line with its nose. To do so means that you are flushing the bird from the angle he expects, thus leaving him master of the situation. This sort of team play between man and dog is, to many hunters, the most exciting part of grouse hunting and, gradually, both members of the team learn more and more about the tricks a modern partridge has in his bag, all of which adds to the enjoyment of grouse hunt- ing and contributes to the hunter's success. A hunter without a dog can still apply such lessons in a general way. If you are in a covert, where you believe grouse to be, size it up before you walk through it: try to anticipate the likely places for a bird to run to before flushing; and try to handle yourself and your comj)anions in such a way that you will disorganize the bird and get a fair shot when it flushes. The authors have observed several old time market hunters, individuals who always hunt without a dog, who can do this to perfection. They are. in a sense, bird dogs them- selves. Finally, from the conser\ation ])oint of view, it should be emphasized that many dead and wounded birds left in the field could be recovered by the more general employment of IV- .iU'. 400 INFLUENCE OF MAN grouse dogs. Even a small spaniel, which is otherwise kept at heel, can very quickly earn his keep by finding a "down bird" or two. Nothing is so disheartening or so contrary to the conservation principle of wise use as to bring down a wily old partridge, only to have to leave him, because he cannot be found, to the skunks, crows and mice, instead of giving him his proper j)lace at the head of a festive board. And then, there is so much to a good grouse dog beyond his utility. One staunch point under a low hemlock, pine or thorn apple, with the sun streaming down through the painted foliage on a crisp October day, makes a picture and provides a thrill which, to use the vernacular, takes the enchanted hunter "right out of this world." And if, perchance, one does his part witli his double gun, and his or her canine highness comes trotting proudly back to deliver the prize — one has something to talk and dream about for the rest of his life. CHAPTER X PARASITISM AND DISEASE By P. P. Levine and Frans C. Goble DISEASE AND ITS CAUSES The So-called "Grouse Disease" — Fa ical Injuries — Chemical Poisons — NutricTORs Which May Cause Disease — Mechan- Parasites — Filterable Viruses tional Deficiencies — Animal Parasites — Plant PARASITISM AND DISEASE IN WILD GROUSE Factors Affecting Different Age Classes — Yearly Variations — Seasonal Vari- ations — Regional Distribution — Relation of Other Animal Diseases to Grouse — Poultry Diseases in Grouse — Parasites of Other Wild Birds — Diseases of Wild Mammals — Relation of Disease to Predation PARASITES OF WILD GROUSE In the Respiratory System — Gape Worm — In the Digestive Tract — Crop Worm — Stomach Worm — Gizzard Worm — Large Roundworm — Large Tapeworm — Small Tapeworm — Cecal Worms — Coccidia — Blood Parasites — Microfilaria External Parasites — Ticks — Louse Fly — Parasites Not Encountered During the Inves- tigation DISEASE IN HAND-REARED GROUSE Quail Disease — Blackhead — Tuberculosis — Aspergillosis — Air-Sac Mites — Perosis THE ROLE OF DISEASE SUMMARY No evidence has been uncovered pointing to a strong causative relationship between disease and periodic fluctuations in grouse abundance, (p. 442). No significant variation in the incidence of parasitism and disease occurred from year to year during the period of the Investigation, (p. 413). No one malady can rightly be termed "the grouse disease". Specific diseases responsible for deaths among grouse in one locality are not found in other parts of the grouse range, while those parasites which have a wide distribution appear largely to be innocuous, (p. 404). 1(12 PARASITISM AND DISEASE IN RUFFED GROUSE Grouse found dead in the field may be the victims of mechanical injuries resulting from shooting, being struck by automobiles, or flying into wires, quite as commonly as from disease. I p. 4041 . The most frequent cause of death from disease in wild grouse in New York State is the inflammation of the saccular stomach due to infection with a small roundworm parasite, Dispharynx spiralis. This affliction occurs oftener in birds from three to nine months old than in any other age class. Most deaths from this parasitic disease occur during the fall and winter. Ip. 411). The stomach worm, Dispharynx, was not found in grouse collected in the Adirondack Moun- tains proper and was infrequently encountered on the periphery of this region although the sow bugs which serve as an intermediate host occur in these areas, (p. 420 I . The most common parasite encountered was the large intestinal roundworm, Ascaridia bonasae, which occurred in over one-fifth of the adult birds. No great numbers, however, were found in any one bird and no deleterious effects could be attributed to its presence, (p. 410). Some parasites, such as the large intestinal roundworm (Ascaridia) , are found throughout the State, while others, the crop worm (Capillaria) for example, are found in limited areas only. (p. 413) . Bird ticks were found to be more abundant in the Adirondack region than in the rest of the State. In that region more birds were infested and more ticks occurred on each para- sitized bird than in other regions of the State, (p. 413). Hand-reared grouse are markedly susceptible to blackhead and '"quail disease"' (ulcerative en- teritis)— the former common to domestic fowl and turkeys. Present grouse propagation methods, however, are effective in keeping these diseases from causing more than occa- sional losses among hand-reared birds, (p. 435). Both the hunter and scientist have become increasingly interested in the healtli of the grouse. The so-called "Grouse Disease" and cycles have been publicized to the extent that they are generally accepted as facts. The study of grouse diseases and their relation to the fluctuating number of the birds in the coverts, as well as the causes and effects of the diseases and how they might be controlled, has been one of the major assignments of the Investigation. Such research is of j)articular importance to the sportsman as it sheds light on many of the question.s paramount to his interest in the grouse. Each year a number of outdoorsmcn discover, for the first time, that their favorite game bird is infected by various parasites. It comes as rather a shock, so they send them to the State Research Center for identification and information. Or they find weak, sick-looking individuals which they fear might have a dread disease. What is the malady, how does it affect their hunting, and what can be done about it? Every nimrod would like to know because, to him. it is of vital concern. The research worker studying disease is equally interested in the distribution and relative k DISEASE AND ITS CAUSES 403 importance of the various diseases to which grouse are subject. The findings here presented are necessarily limited in scope by time and resources. While progress has been made, much is left for future study before all the important questions can be answered. Realizing the necessity for this research, the Investigation's study of disease was initiated in 1931 and has been continued to date, thus permitting observations over a period of eleven years. During that time, 1,728 adult grouse and 1,119 chicks have been collected for autopsy. The birds were taken from all parts of the State, 50 of the 55 counties outside of New York City and Long Island being represented. Table 62 illustrates the distribution of the collections by regions and years. The regions here considered, Adirondack. Catskill and Rest of State, are the same as is used in other chapters of the Report (figure 13 I. TABLE 62. THE NUMBER OF ADULT GROUSE AND OF GROUSE CHICKS COLLECTED FOR PATHOLOGICAL EXAMINATION BY YEARS AND REGIONS Region Year Adirondack Catskill Rest of State Adults Chicks AdulU Chicks AdulU Chicks 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 15 25 57 76 8i 42 30 59 66 59 0 7 50 41 58 32 35 67 40 32 10 17 33 50 56 31 24 45 43 39 0 0 32 19 40 18 21 17 23 21 25 42 93 126 141 73 54 105 110 >)8 0 2 81 62 99 48 59 88 67 60 Total 513 362 348 191 867 .■>()6 Three separate pathologists examined the material collected during their respective assign- ments. This fact, together with changes in the methods and techniques employed as experi- ence accumulated, occasionally raise difficulty in evaluating results. Therefore, in carrying out certain comparisons, it was considered advisable to limit analyses to records of autopsies made by the present authors or under their supervision. DISEASE AND ITS CAUSES It is definitely known that most birds and mammals play host to many plant and animal parasites. It is not equally understood, however, that only a few of these really damage the animals in which they live. \^Tien they do, it is at their own risk. In effect, these disease- producing agents may actually be committing suicide if their activities seriously impair the hosts' well-being. It is erroneous to apply the term "disease"" to the presence of parasites which are causing no alteration in the function or structure of their host. It should be borne in mind that disease is not an entity which enters the body from without but is a process of abnormal activity of certain bodily functions in response to an injury. This process mav or may not be associated with ap])reciable changes in body structure. However, 404 PARASITISM AND DISEASE IN RUFFED GROUSE much of our knowledge of disease is gained by interpretations of the changes occurring in the various tissues and aflected organs. The So-Called "Grouse Disease" In the study of birds whose fluctuations in numbers have so long been observed and specu- lated upon, it is probably natural that some investigators have had preconceived notions about the relation of disease to these periods of scarcity and abundance. The concept and term "grouse disease" came into use as early as 1838 in connection with the fluctuating numbers of red grouse in Britain. Naturally, it was applied in America as interest grew in the study of ruffed grouse. Naturalists, sportsmen, and other amateur observers reported as "grouse disease" their discoveries of louse flies, ticks, "abdominal parasites", fowl cholera and tuberculosis. A professor of bacteriology unhesitatingly remarked that the louse fly was probably the carrier of the "germs" of blackhead in grouse. An ornithologist sug- gested that a small stomach worm might be the cause of "grouse disease" in this country and that the presence of this worm might be associated with the phenomenon known as "crazv flight". Investigators of the British Committee of Inquiry on Grouse Disease" claimed that "grouse disease" in England was mainly attributable to two types of parasitism. One involved a thread-like worm, sometimes occurring in the intestine by the thousands. The other was due to microscopic one-celled animals known as coccidia. These live and reproduce in the cells lining the intestine. In this country, although about 50 parasites and a number of disease conditions have been observed in ruffed grouse, none have been conclusively associated with any widespread de- crease in the numbers of these birds over their entire range. Some parasites that are known to reach a stage of infection serious enough to cause death in New York have not been found in Minnesota or even Vermont and New Hampshire. Others, reported as important in Ontario, have not been encountered in New York. Patho- logical conditions observed in captive grouse are observed in the wild very infrequentlv. No disease recorded to date seems, therefore, to make a serious bid for the title of the "grouse disease" in America or to occupy a position comparable to the parasitisms observed in Brit- ish grouse. Factors Which May Cause Disease One of the convenient approaches to the study of disease is based on the natural classifica- tion of the factors whose presence may excite pathological processes. Some of these may be inherent in the animal itself through inherited traits and tendencies. Others originate in the external environment and, under certain (dnditiniis, become iiiiinicable to the health of the bird. Part of this latter group are physico-chcinical factors such as mechanical injuries, chem- ical poisons and nutritional deficiencies, while part are biological, as rc|)resented by animal j)arasites, plant parasites and filterable viruses. It is necessary to consider each of these in some detail in order to understand their roles in liic lif<" and death of grouse. Mechdiiiiiil Injuries The conditions which residl from physical \iolciicc In llie bodies of llic birds inc known as mcilKiiiical injuries or trauma. Some grouse arc struck by automobiles and trains, others fly into houses or telephone wires. Shooting and predation also come in this category. DISEASE AND ITS CAUSES 405 Although a few cases have been observed in which recovery has been made from minor in- juries, in most cases death ensues sooner or later as a resuh of the structural and functional changes produced. Occasionally shooting or other trauma will result in the perforation of the intestine without serious injury to blood vessels or other vital organs. Death from peritonitis results sometime later. Observations on captive birds have shown that even comparatively minor injuries may cause a disinclination to eat and that death from malnutrition may result a week or so follow- ing injury. As small an injury as the loss of a toenail may lead to severe bleeding and death. On the other hand, birds so severely scalped that their skulls are laid bare, may recover. Chemical Poisons Contacts with the chemical poisons which have come to be of importance in connection with other species of wildlife are of rare occurrence in grouse. During this Investigation one grouse chick, collected for routine examination, presented pathological findings indicative of lead poi- soning, the result of eating and harboring one small lead pellet in its gizzard. The conditions associated with grouse hunting as well as the habits of the birds would logically minimize the occurrence of this disorder. No other instances of either chemical or organic poisoning have been observed. That grouse are remarkably resistant to some of the poisons commonly used in insect or rodent control seems to be indicated by the experiments reported in the chapter on Food Habits fp. 195). Nutritional Deficiencies For the most part, Nature has provided for the adequate nutrition of wildlife in its natural habitat. It is only when broad natural changes or man's activity disturbs the habitat that diffi- culties due to inadequate nutrition are likely to occur. No evidence of malnutrition has been encountered in wild grouse during the course of the Investigation except in birds which were suffering from mechanical injuries or parasitism which interfered with their normal feeding activities. On the other hand, at least one abnormality due to inadequate diet has been produced in grouse reared in captivity. This condition, known as perosis, slipped tendon or big hock disease, has been encountered in young grouse chicks. Animal Parasites The group of organisms most frequently observed by sportsmen and others interested in grouse are the animal parasites. As the hunter picks up a freshly shot bird, he may see among the feathers, winged insects with flattened bodies which scurry about and soon leave the dead game. If he observes more closely, ticks of various sizes may be apparent, closely attached to the skin. Later, when he cleans the bird, he may discover thick white worms a couple of inches long in the body cavity. Or. if the gizzard lining is stripped off preparatory to cooking, thin pink worms may be found lying between the lining and the muscle. These are the comparatively harmless grouse dwellers that are commonly seen. Other animal parasites live in the blood, some actually within the red blood cells, others in the cells lining the intestine. Still others are small flat worms not readily observed with the naked eye. 406 PARASITISM A\'D DISEASE l\ RITE ED GROUSE INFESTATION OF THE GIZZARD WORM (Cheilospirura spinosa) UNDER GIZZARD LINING — A RELATIVELY HARMLESS PARASITE OF GROUSE A NOHMM. I'H()\ INTRICl l.US IKU.IITl COMPARED WITH ONE INFECTED WITH THE STOM \< u woioi I Dispharyitx spiralis) — A RELAIIV i:i.> HARMFUL PARASITE OF GROUSE DISEASE AND ITS CAUSES 407 Animal parasites may damage their hosts in a number of ways. Tapeworms absorb food from the intestine which might otherwise be utihzed by the bird in which they live. Other worms suck blood. Waste products of some parasites are poisonous. In the Investigation of "grouse disease" in England it was observed that injuries produced by parasitic worms pro- vided portals of entry for bacterial infection. I 4 / Sowbug y eaten by gtouse Worm laivae develop in Sowbug Stomach ■worm eg§s passed in droppings of oiouse Adult Stomach Wojtti # \ Eggs hatch \ in % Sowbug — r Egg eaten / Sowbug FIGURE 31. TYPICAL LIFE CYCLE OF PARASITE INVOLVING INDIRECT MODE OF TRANSMISSION — STOMACH WORM ( Displiaryiix spiralis) That the attachmcnl and jiresence of some parasites may cause extensive tissue destruction and indammation has ]ic<-n repeatedly demonstrated during the Investigation by numerous cases of infection by the stomach worm ( Dispharynx spiralis) of which the following is an example. On December 12. 1940. a female grouse about six months old was found in the field in Chenango County. This bird weighed only 292 grams (about 10' ^ ounces). Post- mortem examination revealed that the saccular stomach or proventriculus was enlarged to between three and four times its normal size. The cells lining the stomach were being sloughed off and the tissues below the lining, where numerous stomach worms were found, had been sti- mulated to excessive growth so that the stomach cavity was nearly filled with a plug of mucus and destroyed stomach tissue. 408 PARASITISM A^D DISEASE IN RUFFED GROUSE Infections with animal parasites are picked up hy firouse in various ways. Eggs of the intestinal worm ( Ascaridia) and cecal worm (Ileterakis) are passed from the infected birds in the droppings, develop to an infective stage on the ground and are accidently picked up by other birds while they are feeding. Eggs of some of the other parasites cannot develop to the stage which can infect grouse without first being taken into the bodv of some other animal, gener- ally a snail, insect, or other invertebrate. Grouse become infected with the gizzard worm (Cheilospirura) by eating infected grasshoppers. Likewise stomach worms (Dispharynx) be- come established from feeding on infected pill bugs, while tapeworms are contracted from infected snails, beetles, slugs, ants and flies. Earthworms may be of importance in the trans- mission of the crop worm (CapiUaria) and the gape worm (Syngamus). In these cases it is obvious that the size and number of infections will be influenced by the numbers of the intermediate hosts and the extent to which they arc used as food. A typical life cycle is illustrated in figure 31. Another mode of infection is by the bite of insects. The one-celled animals which cause malaria-like diseases of birds may be carried by black flics and louse flies. Table 63 presents a list of the animal parasites reported from ruffed grouse. Plant Parasites. The plants which are commonlv thought of as parasitic in animals are the bacteria or "germs". These are one-celled organisms, some of which can live in various situations outside of animal bodies for long periods and yet under some circumstances cause disease when taken into the body. Others can live but a short time away from flesh and blood. These must depend on rather direct contact for their transmission. Still others may be normally found existing harmlessly in some parts of the body but are capable of producing serious effects when some other factor alters the function of the organ in which they dwell. In domestic birds bacteria are the cause of many diseases. Some, such as tuberculosis, are occasicmally encountered in hand-reared grouse. However, the plant parasite which has been identified most frequently as a cause of disease in wild grouse is a mold fAsperpilliis fumi- gatus). TTiese microscopic plants are related to the bacteria but have a more complex struc- ture and reproductive process. Filterable Viruses The agents causing cptlain diseases are so small that they can not be observed with ordi- nary microscopes. One of their characteristics is the ability to pass through porcelain filters which arrest the passage of organisms as large as bacteria. Because of this thev arc known as filterable viruses. A disease of horses, corresponding to sleeping sickness in humans, equine encephalomyel- itis, is caused by organisms of this tvpc. Tliis affliction also attacks humans and has been reported from pheasants in the Easf"" and the prairie chicken in ihc \^'cst''. Recognition of such wild reservoirs of disease is of great importance to puiilic health autborilics in the control of diseases of Inimans ;umI domestic animals. Bird ])ox is an important di-casc of this group which is generally seen in domestic poultry and game farm liirds. but it lias also been reported twice from ruffed grouse in New England"'"'^. DISEASE AND ITS CAUSES 409 TABLE 63. THE ANIMAL PARASITES OF THE RUFFED GROUSE AND THEIR REPORTED DISTRIBUTION Parasite Where found Distribution One-celled Animals (Protozoa) Coccidia Flagellates Eimeria dispersa Eimeria aogusta Eimeria bonasae Trichomonas bonasae Cyathosoma striatum Ptychostoma bonasae Histomonas meleagris Trypanosoma gallinarum Small intestine Cecum Cecum Intestine Cecum Cecum Cecum and liver HIta. New Hampshire. Ontario (?) Minnesota Roundworms (Nematoda) Gape worm Crop worm Stomach worm Gizzard worm Intestinal worm Intestinal worm Eye worm Syngamus trachea Capillaria annulata Dispharynx spiralis Cheilospiruru spitiosa Ascaridia bonasae Contracaecum sp. Oxyspirura jwtrowi AprfK-tella sttxldardi PhysalopU'ra sp. Microfilaria Heterakis bonasae Subulura sp. Tetrameres amerirana Windpipe Crop and gullet Saccular stomach Under gizzard lining Small intestine Small intestine Under nictitating membrane In btxly cavity Cysts on muscles In bliHKl plasma Cecum Cecum Saccular stomach New York New York. Southern New England, Michigan New York. Southern New England, Michigan New York, New Eiieland, Michigan. Minnesota, Washington New Y'ork to Washington • Michigan. Minnesota Southern New England .Minnesota New York, Ontario New Y'ork, Pennsylvania. MinnejM)ta New York. Minnesota Strawberry worm .... Experimental infection Mites and Ticks (Acarina) Megninia sp. Trombicula microti Laelaptinae Liponyssus sylvarium Cnemidocoptes mutans Gytoleichus nudus Haemaphysalis leporis- palustris Haemaphysalis chordeilis Attached to skin or feathers Attached to skin or feathers Attached to skin or feathers Attached to skin or feathers Attached to skin or feathers In air sac Attachel>lici4 to tlic more sedentary, wingless ectnpnrasitic relatives of the spiders and mites. One might conclude from the table thai intestinal wtirms. which occur in such a high per- centage of the birds, arc of the greatest importance. This \ iew. however, can not be siil>staiili- ated when the ability of the parasite to produce disease is considered. The inlestinal wDrrn ( Ascariilia honasdc) . on account of its size ami fretpiencN of occurrence, is the one most com- monly obserxcd by si)ortsmen and which the\ arc iikcK to regard as a serious enemy of grouse. The organism which is, however, most capable of producing disease is the seldom seen stomach worm which occii|iies luil fourth place in frctiiicniy of occurrence in chicks and second place in adults. PARASITISM AND DISEASE IN WILD GROUSE 411 It is likewise important to point out that the number of parasites found in any one bird does not necessarily indicate the importance of the infection, since the presence of as many as 78 intestinal roundworms has been noted without evidence of damage to the host. On the other hand, only a few stomach worms may be found in a badly diseased stomach. Table 65 shows the average numbers of parasites per bird of those infected and the maximum number encountered in any one based on observations made during the Investigation. I'ABLE 65. AVERAGE AND MAXIMUM NUMBERS OF PVRASITES ENCOUNTERED IN GROUSE IN NEW YORK— 1931-19H Parasites Average number Maximum number Internal: 20 6 5 2 28 100 Crop worm 20 45 Intcstiniil worm Gape worm 78 8 External: 'I'icks . 300 i.oiise nies 10 Since this lack of relationship exists between the abundance of parasites and their pathogen- icity, a tubulation of the disease conditions encountered offers information of greater signifi- cance in the evaluation of the role of these itnaders. TAHLh; ()(.. RELATIVE OCCURRENCE OF I'ATIIOLOGICM, CONDITIONS IN ADULT AND YOUNG (iROUSE IN NEW YORK— l'):il-l<)U Age I'athological CDlldilioii Adulu Cliiclis Per cent of incidence Per cent of incidence Stoiiiiicli inlliininwition. . . . Crop iiilUitiiniation Mold infection 10.9 l.O 0.5 O.t (I.I 0.1 0.1 o.i' 2.7 Intestinal inllannnatlon . . . Gizzard inllammation 0.6 Egghoinid Lead ])oisoning d i 0.1 Tick infestation An examination of table 66 reveals the significance of stomach worm infection in its true light, since it is apparent that inflannnatory reactions due to this helminth lead the list of disease conditions observed. \S ith the exception of one case of tick infestation, all animal parasitism resulting in death has been attributable to this small roundworm. The only other fatal infections observed were due to aspergillosis. This disease, caused by tile invasion of the bodv bv a mold or fungus organism, was the onlv maladv which was found 412 PARASITISM AND DISEASE IN RUFFED GROUSE ill both wild and captive grouse. In the former it was encountered only a few times and in widely separated parts of the State. Inflammation of the crop, intestine and gizzard, associated with infections by crop worms, tapeworms and gizzard worms respectively, were not of a serious nature and no deleterious effects on the well-being of the birds were observed. Certain variations due to geographical, seasonal and age differences have become apparent during the course of the Investigation. These indicated that a table of incidence based on total collections presented information of an extremely general nature. To take but one ex- ample, the incidence of the stomach worm ( Dis pharynx) in all birds over three months old is listed at 12.7 per cent. Subsequent analysis revealed that this parasite was almost entirely absent from the .Adirondack region, appearing only occasionally on the periphery. Further examination of the data showed that 29.1 per cent of the birds of the year (from three to nine months old) were infected, but adults taken during the spring and summer had infections in only 8.7 per cent. Accordingly, in order to complete the picture, the variations due to age of the birds, seasons and regional distribution are discussed in the following sections. Factors Affecting Different Ace Classes That certain organisms commonly parasitize adults while others are more liable to be found in young hosts, has been observed frequently during the course of recent pathological examin- ations. This phenomenon is clear-cut with reference to certain of the parasites encountered in grouse but is less well defined for others. Incidences of the commoner parasites in the differ- ent age classes are shown in table 67. TABLE ()7. INCIDENCE OF THE COMMON PARASITES OF GROUSE IN NEW YORK BV AGE CLASSES— 1931-19 a Per cent of incidence of parasites* Agc! classes Total number cxaroiiied Ta|»ewonnsA Coccidia Intestinal worms Stomach wormst Gizzard worms Tickst 0 to 1 months 270 267 117 189 332 172 16.6 32.2 2.0 1.6 9.9 4.6 12.2 1L<) 1.1 1.1 0.6 0.0 0.0 10..5 2«.6 1<».6 2.5.9 16.9 0.0 8.2 23.8 29.1 11.7 1 l.ll 0.0 ^8 6.1 11.1 1(1.. 3 9.6 17.9 2 to .'J iiumt lis .... LI Spriii).' 1111(1 siiiiimiT Mllulls 11.6 Fill! Mild winter * Thp romliineil inculetifr of inlrrnal parasites indicated here fur iho three age claniie* cuverine the chick period (under 3 nioritlia) diflern iili|;Iili\ from ihr ligiirrft in tab If 6i hcL-ausc rrtain specimens included there, although they were known to iall witliin this run^c i-ut very soon there- after ta|»e\v(>rms and coccidia are found. Botli of these are relatively al)undant in June and July l>ut infrequently found by August. Most of the tapeworms found in these young birds are a large form (Raillietina tetragona) also found in domestic poultry. It is presumed that PARASITISM AND DISEASE IN WILD GROUSE 413 the infection is gained by eating ants, which early in the summer form a large part of the animal food of the chicks. Coccidia, which have a direct life cycle, are obtained by accidental ingestion of the infective stages during feeding. Neither coccidia nor the above tapeworms are found in any appreciable number of adults. Ticks, which occur on about 12 per cent of both chicks and adults during the summer, are most frequent in July. During July, the three parasites which occur most frequently in adults begin to make their appearance in the chicks. These are the stomach worm (Dispharyii.x spiralis) gizzard worm I Cheilospirura spiuosa) and intestinal worm ( Ascaridia bonasae). In the birds of the year during the fall and winter, the stomach worm occupies a prominent position. Although it seems capable of producing lesions equally serious in adults as in young birds, the relative incidences probably indicate its greater importance in the latter. Yearly Variations In view of the numerous reports on the cyclical nature of grouse abundance and scarcity, the analysis of the data on parasitism on a yearly basis was regarded as one of the most im- portant phases of the Investigation. Although, in some instances, superficial inspection of the data seemed to indicate that yearly variation occurred, this was found to be due largely to changes in the intensity of collecting in localities or seasons in which the parasite might be more or less abundant than the average. Since these seasonal and regional effects had to be removed in treating the material, the in- cidence of only the numerously occurring parasites could be compared from year to year. These analyses were carried out for intestinal roundworms, stomach worms, gizzard worms and tapeworms in adults and for tapeworms, intestinal roundworms, stomach worms and ticks in chicks. No evidence could be adduced to support the idea that significant changes in the incidence of parasites took place from year to year during the course of the Investigation. It must be remembered, however, that no substantial decline in the grouse population occurred during this period. Seasonal Variations The general view that disease and parasitism is more prevalent during the warmer months is borne out by observations on tapeworms and intestinal worms in grouse, but it is not appa- rent from the data on stomach worms. No explanation for this difference is suggested by the data or the facts which are known concerning the parasites and their method of dissemina- tion. The drop in incidence of coccidia from 12 per cent in July chicks to less than two per cent in fall juveniles is probably attributable to age and iinniunity rather than season since the incidence in summer adults is also very low. Regional Distribution Analysis of the data on parasites taken during the Investigation reveals three salient exam- ples of regional distribution of the parasites observed. A higher incidence of ticks was found in the Adirondack region. In the same region, an exceedingly low incidence of stomach worm was noted. Autopsies revealed no records of crop worms outside of the Hudson Valley. With the exception of these three instances, the distribution of the parasites was statewide. It should be pointed out here that the figures used in comparing the tick populations on "rouse in the various regions of the State have been based only on those specimens taken by 414 PARASITISM AND DISEASE IN RUFFED GROUSE special collectors during the siummcr iiKuiths. These birds were placed in cellophane sacks im- mediately after shooting so that ticks would not escape in transit. They were found on about 26 per cent of 190 grouse chicks from the Adirondack region and ai)out 6 per cent of 510 from the Catskill region and the rest of the Slate. The number of ticks on a grouse was also higher in the Adirondacks. averaging 38 there and five in the other regions. The greatest number found on Adirondack birds was 300 while in other regions 26 was the maximum. TMKKK KXIKHWI. l>\li\sn'KS f)!' TlIK RUFFED GROUSE AND THE SOWBIC. INTERMEDIATE HOST OK THE ST()\I\(TI WORM f Dls/iliai vrix ) Sntvhii/; ( I'orrellio srabrr) Louse Fh < Lynchia americana) HirtI I.tiiise I I.ngnpoerus perplr.xiis) Bird Tick I Haemaphy.salis chonlrilis) PARASITISM AND DISEASE IN WILD GROUSE 415 Examination of over 300 Adirondack region grouse which were old enough to show stomach worms, revealed the presence of this parasite in only four cases. All four of these birds were collected on the periphery, two on the northwest boundary adjoining the St. Lawrence valley, and two on the southeast boundary adjoining the Hudson and Lake Champlain watersheds. None was found in the Adirondack interior. It has been reported that infections with this worm are contracted by eating sow bugs. Specimens of the latter (Porcellio scaber) have been found in the crop contents of grouse chicks from this region, so it would appear that it is not the absence of this intermediate host which accounts for the negative findings. The croj) worm was found only in the counties bordering on the Hudson River. Allen and Gross" reported it oidy from New F^tigland and Columbia County. New York, while Cram^ reported that the infection of quail with these worms was quite localized. Sinte this is one of the parasites which has also been reported from poultry, it is of interest to note that it is seldom encountered in chickens in New York. Beaudette" reported only two cases in almost ten thousand autopsies of New Jersey poultry. Relation of Other Animal Diseases to Grouse Speculation c cniulilioned consid- erably by factors concerning relative tick populations on grouse and cottontails and the species of ticks involved. In Minnesota where tularemia lias been reported as naturally occurring in game birds, it is common to find several hundred licks on a grouse, cottontail or varying hare, while in New York the average and the maximum infestations are much lower. Cottontails in May have been found to carry from one to 184 licks I average U)). The maximum number of ticks found on grouse in the summer was 300 in ihe Adirondacks (average 38) and 26 in the rest of the State (average 5). A large percentage of the ticks found on grouse in New York are the bird ticks (H. chordcilis). Up to 87 rabbit licks have been found on varying hares. PARASITES OF WILD GROUSE 417 Although Green and Shillinger"' mention only the genus Haemaphy salts with the implication that the rabbit tick is the only one involved with the transmission of tularemia to grouse, Parker, Philip and Davis""" present evidence pointing to the probability that the bird tick (H. ch-ordeilis) acted as a vector in an outbreak of this disease on sage grouse. The rarity of tularemia in New York may be indicated by the fact that, during the decade 1932 through 1941, there were only eight cases of tularemia among persons which could be traced definitely to animals having their origin within the State. Of these, the species impli- cated were as follows: cottontails — 4, muskrats — 2, fox — 1, deerfly — 1. In two of the cases where the infection was suspected of having been contracted from cottontails, a collector from the Conservation Department immediately went into the field and trapped or shot all the rabbits which could be contacted in the vicinity in which the original suspect was taken. Blood sam- ples from these were subjected to agglutination tests by the Department of Public Health and the animals were sent to the Research Center for post-mortem examination. F'.ighty-seven rab- bits were taken from one locality and 27 from the other. In all instances blood tests and autopsy findings were negative for tularemia. It seems probable that further observations on the inter-relation of tularemia, wild birds and mammals, and ticks will come from areas where tularemia is more prevalent. Whether the presence on grouse of certain miles, ordinarily found on small rcidents (Laelap- tinae, Trombicula microti), is accidental or significant is a matter for speculation. It has been suggested that rodent mites related to these may be involved in the dissemination of tularemia among meadow mice"'". Relation of Disease to Predation The relation of predators to the elimination of diseased animals has been the subject of many discussions. During the course of the Investigation, a number of grouse were received which had been killed by predatory mammals or birds. Of these, 33 were satisfactory for post-mortem examination. In none of these was parasitism or disease present to a degree which suggested that either might be a factor in the susceptibility of the bird to predation. Nineteen cases were completely negative and the parasitism in the others differed neither in incidence nor degree from that found in a normal population. We have, however, a number of records of birds which were weakened by a result of stom- ach worm infection and were readily shot or picked up by dogs. It can be inferred that these diseased grouse would also have been available to other predators. PARASITES OF WILD GROUSE In the Respiratory System Gape worms were the only animal parasites encountered in the respiratory tracts of wild ruffed grouse during the Investigation. The occurrence of the fungus infection, aspergillosis, in both wild and hand-reared birds has already been mentioned. Discussion of this disorder, which is more likely to be found in captive grouse than in the wild, has been relegated to the section on disease in hand-reared birds. Air-sac mites, also discussed later, were found only in captive grouse in association with poultry. Gape Worm (Syngamus trachea) Distribution. This parasite is found in most parts of the world. It has been re- 418 PARASITISM AND DISEASE IN RUFFED GROUSE ported ill chickens, bobwhitc quail, turkeys, pheasants, peafowl, red grouse, sparrows, thrushes, crows and a number of other birds as well as in ruffed grouse. It has been considered as asso- ciated with pheasants and domestic birds for so long that it was surprising to find it occurring in areas in the deep Adiroiidacks where poultry has never been raised. It has been found in several widely separated localities in New York but its incidence is so low that its absence in specimens from other regions is no indication that it is non-existent there. Description. This worm, sometimes called the "Y" worm, because the male and the female are joined together in the form of a Y, is found attached to the mucous membrane of the windpipe. The worms are blood-red in color. The males measure up to ' i incli long, and the females up to 'J^ inch in length. Cram"" has given a complete description. Life Cycle and Dissemination. The eggs are deposited in the windpipe by the female, are coughed up, swallowed and passed out with the feces. Under optimum conditions of tempera- ture and moisture, development to the infective stage takes |)lace in about seven (ia\s. Infec- tion results when a susceptible bird eats either the embryonated eggs or certain invertebrates, such as earthworms and snails, in which the infective larvae are loosely encysted"-*". Within a day after ingestion, the larvae migrate to the lungs. According to W'ehr'" this migration takes place by way of the blood stream. The males and females jjair as young adults in the lung and then move up the bronchi to the trachea where attachment occurs. The worms ma- lure within 1 7 to 20 days after infection. The spread and maintenance of this infection in any one species is influenced by the follow- ing considerations: 1. Gape worm larvae encysted in earthworms mav remain viable and produce infection for longer than four years as demonstrated by Taylor*". 2. Man> species of small wild birds are found to be infected with the same species of worm. .■^. In some species of gallinaceous birds there is a definite age resistance to infection. Adult chickens, for instance, are rarely infected, while very young fowl become infected with comparative ease. I iirkc\s on the other hand retain the infection at all ages. It is in- teresting to note that the incidence of gape worm infection in grouse chicks (0.0 per- cent) is more lluiii four times the incidence in adults. It is possible that the extreincK low incidence of the iiifc( tion in \ew York groii>c i> due to the fact that the ex|)osure to infection may be very light and scattered. Gape worm infection in domestic chickens and turkeys in this State is seldom encountered by the diagnostic laboratory at the New York Slate Veterinary College. Serious reservoirs of infection may exist on pheasant farms. Here the gape worm disease has been known to lake a lieavv toll of pheasant chicks unless the young birds are raised on soil on which pheasants have not hccii prc\iip|isl\ reared for a period of at least three \ears. Pathonenieity. The abilitv of ga])e worms to produce disease in pheasants and domestic poultry is well known. The blood .-iucking activities of the worm and the nu'chanical block- ing of the windpipe are serious enough to cause the death of heavily infected birds. In native wild grouse, however, the number of parasites encountered has never been more than eight pair and in none of these birds was there evidence of serious injury. The onl\ evidence that S. trachea can be serious for grouse was observed in 1932. A wild trapped female grouse shipped to New York was found to have died as the result of suffocation by 35 worms packed in the lower end of the windpipe. This bird was being used as a breeder and was confined in a large pen on ground where nincs are likewise susceptible. In New York, the birds found to be infected with this worm were all taken in the Hudson River Valley. It was nol fomul in cilln r |iarl> of llir Skili-. Description. The crop worm is a long, thread-like form that is frequeiitb found imbedded in the lining of the crop and esophagus of wilfl grouse. The niaics measure up to I inch long and the females up to 2Vi inches. Though it is fairly long, it is so narrow that it is very difTicult to observe. IJfe Cycle and Dissemination. Wehr'™ has found that earthworms act as intermediate hosts. Apparently the larvae undergo a period of develoi)mcnl in the earthworm iiefore infection of the avian hosi utii>M does not extend into the coniferous forest regions or so-called Canadian Zone. The reasons for this are not known. Descrijilion. The sloniacb worm. Displiarynx sjiiralis. is a short, fairly thick, white, coiled nematode (roundworm) that is found deeply imbedded in the lining of the proyentriculus (saccular or glandular stomach). The males are about \'\ inch, the females about % inch in length. /-;/(• Cyrlr anil Dissemination. The life cycle of this worm was worked out by Cram'". It was found that two species of land crustaceans, the common sow bugs or pill bugs, Porcellio PARASITES OF WILD GROUSE 421 scaber and Armadillidium vulgare, acted as intermediate hosts. The eggs laid by the female worms pass out with the feces and are ingested by the sow bugs. After a period of develop- ment of about 26 days, the larvae become infective. When infected sow bugs are eaten by sus- ceptible birds, the larvae are liberated and invade the lining of the stomach. The worms reach maturity in 27 days. It is not known how long the eggs of this worm remain viable or its larvae live in the in- termediate host as a source of infection. It is significant that, in practically every instance, grouse found to have severe lesions or to have died as a result of infection, were encountered during the late fall and early winter. In fact, the incidence of the infection was highest during that time of the year. Apparently the birds pick up stomach worms during the summer and fall. The effects of the parasitism are seldom evidenced until the worms have matured. The smallest bird in which Dispharynx was found was a young male which weighed 144 grams (about S oz.) collected on July 27. The earliest record of occurrence was in a young female collected on July 20. That she was in good flesh may be judged from her weight of 292 grams (about 10^2 oz.). at the time of collection. That infections with this worm may be quite localized was illustrated by collections made on Connecticut Hill in 1934. Five birds out of 17 in one covert were infected while no stomach worms were found in 13 birds taken in another covert 200 yards away. Each covert included about 250 acres. They were separated by open land. PalliDficiiirilv. Dispharynx s/iiralis is |)robab!y the most ])athogenic parasite of wild grouse in the more settled parts of the Northeast. Heavy infections with this worm result in a severe inflammation of the glandular stomach or prevent riculus. This becomes filled with a thick, white, shiny mucus. The proventriiular wall becomes tremendously thickened, sometimes to three and four times its normal size. There is an extensive destruction of the glandular tissue and of the muscular layers of this organ. This tissue destruction may progress to such an extent that perforation of the glandular stomach may bring on peritonitis. Tissue necrosis quite often accompanies this parasitism. In many instances the proventri- culus, due to destruction of its musculature, becomes quite flaccid and may telescope in its entirety into the gizzard. Cases have been found where healing of the lesions resulted in the production of masses of dense connective tissues resulting in a partial or complete closure of the organ. Large numbers of worms may not always be found in spite of extensive tissue damage and sometimes none are encountered. Undoubtedly many of the parasites pass out of the bird with the sloughed off tissue or they may become detached during the time the bird is in a dying state. On the other hand, 100 or more of the worms have been dissected out of a single bird. Allen and Gross" have reported as many as 228 in one stomach. The invasion of the proventriculus by the stomach worm sometimes results in emaciation, weakness and finally death. Hunters occasionally report that birds, later found to be seriously infected, were sluggish in flight or that their dogs, unaided, ran down and caught the birds. This is undoubtedlv due to weakness resulting from loss of function of the proventriculus through tissue destruction and plugging of the stomach cavity. Light infections with this parasite have been encountered where no apparent ill effects on the birds were noticeable. In these cases, however, the parasites were few in number and only a small portion of the proventriculus was involved. 422 PARASITISM AND DISEASE IN RUFFED GROUSE Although the stomach worm was found to be the most pathogenic species in ruffed grouse, in no sense should it he iinafrined that infection is alwavs fatal. Considerahle indirect evidence was gathered indicating that hirds may lose their parasites. In a number of instances healed lesions were found in the proventriculus, characterized by extensive scar tissue. Since no worms could he found in these birds, it is safe to interpret these findings as instances of recovery from the parasitism. One case is worth mentioning here since it represented a rather unusual situation. Fifty- three stomach worms were removed from a markedly enlarged proventriculus. Yet the bird, a male, was in excellent condition for it weighed 622 grams (about 22 oz.). It is quite likely that this infection may have been recently acquired and the deleterious effects of the ])arasi- tism may not ha\e set in. The extent of the infection and the resistance of the host undoubt- edly play a major role in determining whether or not the parasitism will evcntnallv kill the bird. With the exception of one serious case of tick infestation, stomach worm infections were the cause of death of all wild grouse without mechanical injuries that were found dead in the field. The relative infrequency with which dead grouse are picked ii|> in the field gives this fact even more significance. During the months of October and November. 19.33, 11 ruffed grouse were referred to the Investigation by the Coiniecticut State Hoard of Fisheries and Game. Tlicse birds had been submitted by hunters and other interested individuals whose attention had been attracted by their abnormal behavior. Seven of this number were found to be heavily infected with the proventricular worm. In all probability similar conditions prevail in other areas where this parasite is found. In no sense, however, should it be assumed that ruffed grouse in all regions suffer from this infection. Attention has been called to Boughton's" survey in Minnesota where not a single instance of infection was encountered though .SOO birds were autopsied. Again. Muellers sur- vey^' of niffcd grouse in New Hampshire indicated the absence of D. spiralis in any of 46 grouse examined. These records clearly indicate the danger of attempting to draw conclusions from data collected in restricted areas. Incidence and Ini /lortance. Aside from being a {)rime factor in killing birds, stomach worm infection undoulilediv adds indirectly to grouse niortalitv. Parasitized birds, especially in the latter stages of the disease, may become so weakened that iheir flight is seriously impaired. There is no doubt that predators may make short work of many such birds, if found. Infection with stomach worms takes place earl\ in the life of the chick but the worms are not ajiparcnt until the second month when 8.2 per cent of the birds are infected. The incidence rapidly rises to 29.1 per cent in the three to nine months ajjc ;^innii and ihcri >lMin|is sliarph to 8.7 per cent in the spring and summer adults, Ihc high prccentage of infection of the young bird.-- ami the low pciccEilagc found in those wild ha\(' survived the winter, raises a very pertinent question as to the reasons for this change of incidence. It is problematical if the parasites are lost during that cold period or whether the infected birds died and only those free of parasitism lived to be examined in the spring and summer. The answer depends upon further research. If it were found llial the stomach worm caused the death of most of the infedi-d \onng prou>c. llic laii;i' nuinliers involved would explain one major source of loss. Since, in llic fall and winter adults, the incidence only rises to 1 I per cent, the further PARASITES OF WILD GROUSE 423 question to be answered is whether this represents an age immunity or one acquired through recovery from previous infection. The solution will require the infection of young birds held in captivity and the following of the subsequent course of the disease. The overall incidence of infection in grouse over three months of age was 12.7 per cent. Despite the fact that the stomach worm occurs half as often as the intestinal worm during the same time period, the former possesses a much greater potentiality for damage to the host. Where it occurs in New York, D. spiralis infection is the most important worm parasite among wild ruffed grouse. The determination of its relative role in grouse population fluc- tuation is not easy. No significant variation in occurrence from year to year and no relation to cyclic diminution can be demonstrated. During the period of the Investigation, no epizootic phase of the disease has been observed. The scope of this survey can only justify the conclu- sion that this parasitic infection may be a cause of grouse mortality in certain areas. Gizzard Worm (Cheilospirura spinosa) Distribution. This parasite has been found in ruffed grouse from Michigan, New York, Minnesota. Pennsylvania, New Jersey, Wisconsin, and Massachusetts, and in bolnvhitc quail in Virginia and Tennessee. In New York it has been found to be widely distrilnitcd through- out all the region. Description. Cheilospirura is a moderately slender pinkish worm found beneath the cor- neus lining of ihe gizzard. The females measure up to T j inches, the males to ^'\ of an inch in length. Cram gives a conii)lete description"". Lije Cycle and Dissemination. Embryonated eggs, discharged in the feces, are ingested by grasshoppers. The worm larvae migrate from the digestive tracts of these insects and become loosely encysted in their bodies. After 25 days, the infective larvae are developed. Wlien susceptible birds eat infected grasshoppers, infection results. Maturity of the parasite within the bird is reached in 45 days. Cross transmission of the parasite from ruffed grouse to quail was proven experimentally. Aside from the fact that the life cycle of this jiarasite has been worked out, very few of the factors that determine the spread and pathogenicity of this worm are known. Cram" noted that with the gizzard worm infection in chickens (C. hamulosa) a certain proportion of the invading nematodes failed to develop. She suggested the possibility that the health of the individual fowl might play a role in explaining this phenomenon. It is likely that a similar condition exists in grouse thus infected. The earliest date on which Cheilospirura was found in grouse chicks was July 15 when it a|)peared in a female weighing 221 grams (almost 8 oz.). Pathogenicity. In experimental transmission experiments, Cram"" demonstrated the patho- genicity of the gizzard worm for ruffed grouse and quail. Hemorrhage and necrosis of the gizzard lining were noted, these changes being accompanied by flabbiness of the gizzard mus- culature. Over a period of 84 days, an unnatural growth of this organ was produced in one quail with a heavy infection. Fisher"" stated that in Michigan, where infection during 1935 and 1936 was over 50 per cent. "Many of the grouse were so heavily infected with these para- sites that part of the lining of the gizzard had been destroyed and there was destruction of the surrounding tissue." On the other hand, Boughton"" failed to find serious lesions in infected grouse in Minnesota. 424 PARASITISM AND DISEASE IN RUFFED GROUSE The heaviest infection observed during this Investifration was 45 worms, the presence of which caused only minor lesions in the gizzard. T^Tiether these parasites would have caused severe damage in time can only be conjectured. Incidence and Importance. Gizzard worms were found in birds from practically all areas of the State. The incidence of infection gradually rose from none in grouse chicks less than a month of age to II. 1 per cent infection in the spring and summer adults. The incidence of parasitism with this worm in grouse of three months or over was 7.8 per cent. This figure covers the total number of birds examined in this survey. There was no significant difference in the incidence of this parasitism from vear to year. Though the pathogenicity of this worm for birds has been demonstrated in certain instances, in the light of observations during the Investigation, gizzard worm infection is considered to be of little consequence in controlling grouse abundance. Large Roundworm (Ascaridia honasae) Distribution. This parasite has been found in ruffed grouse in Maine. Michigan. Minne- sota, Wisconsin, Pennsylvania. Massachusetts, and has been found to be widely distributed throughout the three regions in New York. Its distribution in other birds is uncertain. Until recently, A. honasae was not distinguished from another form. .4. lineala. which is common in poultry and has been reported in various species of game birds. Description. Ascaridia honasae is a fairly large, whitish, roundworm commonly found in the small intestine of wild grouse. The males measure up to 1% inches long and the females up to 2 inches. A complete description is given by Wehr'"'. Quite often these worms are found within the body cavity of grouse that have been shot, for perforation of the intestine permits the parasites to escape. Life Cycle and Dissemination. AckertV description of the life cycle of A. lineata of the chicken is summarized below. It is probable that the life cycle of A. honasae is quite similar. In the intestine of the bird, the female worms lay eggs which are eliminated with the feces. Under suitable conditions of temperature and moisture, enibrvonalion takes place in from 12 to 16 days. After the birds take in these embrvonated eggs with the feed and water, the young larvae are liberated in the forepart of the small intestine. During the first nine days the larvae live in the intestinal interior. During the following week, development and growth proceed within the tissues of the gut. At the end of this time the worms emerge from the intestinal wall and remain in the interior until maturity is reached. This occupies about .SO davs. Ackert and Herrick" and Levine'^ have noted that in chickens there is a definite age imniiiMity to A. Hneata. Furthermore it has also been established that not all of the young worms that emerge into the inside of the intestine succeed in establishing tbcmsclvcs. The viabilitv of the eggs of A. lineata has also been investigated by Ackert and Cautlien". and Lcvinc'^. In gen- eral, drying and exposure to the sun has a marked lethal effect on the ova. while weathering in the shade tends to greatly prolong the period of survival. I,evine"° showed that the length of time the eggs remain viable when exposed to the cold de- ])cnds on whether or not thev are embrvonated. Kinbr\onatfd ova were destroyed after two weeks of freezing while non-embryonated ova survived a severe New York winter and devel- oped the following spring. It is safe to sav that pmbablv all of these factors plnv a part in infections of wild grouse. Pathogenicity. The harmful effect of A. lineata infection in young chickens has been des- SOME INTERNAL PARASITES OF THE RUFFED GROUSE Gape Worm (Syngamus trachea) Stomach ff'orm (Dispharynx spiralis) Intestinal Worm (Ascaridia bonasae) Crop Worvi ( Capillaria annulata) Gizzard Worm (Cheilospirura spinosa) Cecal Worm (Heterakis bonasae) 426 PARASITISM AND DISEASE IN RUFFED GROUSE cribed by Ackert and Herrick'. They found that definite symptoms of parasitism and death resuhed from heavy infections. The greatest injury to the birds took place during the 10th to 17th day of the infection when extensive tissue invasion and destruction was occurring. Although A. bonasae in grouse was found more frequently than any other nematode, no in- stances of the pathogenic effects of this worm were observed. The worms were nearly always adult and occurred for the most part in small numbers, five being the average number found. Incidence and Importance. Grouse chicks pick up the infection when quite voung. 10.5 per cent of them becoming infected by the second month after hatching. The following month shows the infection rate more than doubled and from that time on there is a fluctuation in the incidence of infection. A higher percentage of grouse is infected during the summer months and a lower percentage infected during the fall and winter, table 67. The data avail- able suggests that Ascaridia infection is unimportant from the standpoint of brood mortality. This large roundworm is one of the most frequently encountered parasites of ruffed grouse in New York State. In its adult stage it feeds on intestinal contents. On account of its size, it is the parasite most often observed and rated as important by amateurs. The number of worms found in each bird is small. In one instance, however. 78 worms were found in the in- testine of a male bird. Despite this heavv infection, no deleterious effects were observed. No significant vearly variation in incidence took place during the course of the Investigation. Insofar as this study is concerned, no evidence was uncovered which would indicate thai in- fection with A. bonasae is a factor in reducing the grouse population in the Northeast. Large Tapeworm ( RaiUietina tetragona) Distribution. This parasite is found in the small intestine, occasionallv in the cecum, of ruffed grouse, chickens, bobwhite quail, turkeys and guinea fowl. It has been reported from Europe and Asia as well as from North and South America. It occurs in all three regions of New York. Description. These tapeworms mav attain a length of 10 inches. Thev appear as creamv white ribbons about ^s of an inch broad, composed of manv segments. Thev possess, as a means of attachment, four suckers as well as a crown of minute hooks. Lije Cycle ami Dissemination. The segments farthest from the attached end of the worms contain many eggs. As these segments become detached from the rest of the parasite and pass out with the droppings, the eggs become available to various invertebrate animals which serve as intermediate hosts. A number of organisms have been reported as the vectors of this common tapeworm of grouse but the most likely suspect in this area is the ant. Horsfall"* reported ants of the genera Tetramorium and Pheidole to be involved in this life cycle. While these particular insects have not been found in the food analyses bv tlic InvcsliLiation. manv of their close relatives are taken by the young chicks. Since infection with this parasite must fullnu the eating of some intermediate host, it is not surprising to find that most of the cases of parasitism with this form occur in the wild grouse chicks, whose early diet contains such high amounts of animal matter. Conversely, in the vege- tarian adult grouse this tapeworm is seldom found and when it does occur it represents the chance ingestion of an infected intermediate host. Pathogenicity. Although experimental work with tapeworms (Davainea proploltina) in poultry"" has furnished definite e\idence of the deleterious effects of infections with these par- i PARASITES OF WILD GROUSE 427 asites on growing chickens, no similar evidence could be adduced from observations on tape- worms in young grouse. In a few cases, excessive mucus production by the intestinal cells was attributed to the presence of the parasites but no more serious changes were observed even in heavy infections. The weights of parasitized birds compared favorably with those not infected but of the same age. Incidence and Importance. Though the incidence in chicks less than two months old is rela- tively high, being 16.6 per cent during the first month and .32.2 per cent in the second, the lack of lesions and the good physical condition of the infected birds seems to indicate that little harm is caused by the presence of this parasite under natural wild conditions. As the summer progresses the number of chicks infected drops off to 2 per cent in August and there- after it is very seldom found. Small Tapeworm ( Hymenolepis microps) Distribution. This parasite is found in the ruffed grouse in iNew York and Ontario, in the ptarmigan, and in the red grouse, the black grouse and in the capercailzie in Europe. It occurs in all three regions of New York. Description. The tapeworms of this species are found attached by their head end (scole- cesj to the wall of the upper portion of the small intestine. To the naked eye the parasites ap- pear like long thin filaments. A detailed description is given by Shipley"". Those found by the Investigation were identified by Jones'" whose account is the first record of this worm in ruffed grouse in this country. Life Cycle and Dissemination. The life cycle of this parasite is unknown. Related tape- worms of the gallinaceous birds reach infectivity in insects, particularly beetles. Pathogenicity. In one heavy infection with this tapeworm a small blood streaked area in the duodenal mucosa was found where a great many scoleces were imbedded. A catarrhal in- flammation of the upper portion of the small intestine was observed in another case where the infection was severe. In practically all the other instances, however, the number of tapeworms in any single in- testine was relatively few and no gross lesions were observed. No evidence was uncovered to establish that infection with tapeworms could kill ruffed grouse. Incidence and Importance. Although this parasite was sometimes encountered in chicks it is regarded as more typical in adults. In juvenile birds from three to nine months old, its inci- dence was 1.6 per cent. Almost 10 per cent of the spring and summer adults were infected, but less than 5 per cent of the fall and winter adults are thus parasitized. This tapeworm can not be considered of importance as a pathogen in wild grouse on the basis of any observations made during the Investigation. Cecal Worms (Heterakis bonasae) Distribution. This roundworm has been found in grouse in all three regions in New York. It is probable that many of the previous records of cecal worms from grouse, designated as Heterakis gallinae. a common parasite of poultry, may actually represent records of grouse cecal worms. It is therefore almost impossible to establish the distribution of this worm in other hosts and in other regions. Description. Heterakis bonasae are small white worms measuring about V2 inch in length. 428 PARASITISM AND DISEASE IN RUFFED GROUSE Life Cycle and Dissemination. The life cycle of the closely related cecal worm of chickens has been worked out and is essentially the same as that of the intestinal roundworm, the ascarid. Eggs of the worm pass out in the feces and develop to the infective stage in about seven to twelve days. Upon ingestion, the worm larvae are liberated and quickly find their way to the ceca where development to sexual maturity is completed in 56 days. Undoubtedly the cecal worm of grouse undergoes a very similar development. Pathogenicity. No evidence of injury to rutled grouse by cecal worms has been uncovered. Usually the number found are very few. It is interesting to note that in chickens and turkeys, the cecal worm egg is capable of trans- mitting blackhead. No case of this disease in wild ruffed grouse has been encountered by the Investigation, though it is occasionally found in grouse reared in captivity*. Incidence and Importance. The Ughtness of tlie infection and the lack of demonstrable damage done by H. bonasae make it a comparatively unimportant parasite of grouse. Only 1.8 per cent of grouse over three months of age were found to be infected. Of the chicks, only 0.1 per cent were infected. Coccidia (Eimeria species) Distribution. One species, Eimeria angusta was described from the ceca of ruffed grouse in Labrador and Alaska'". It has also been found in the spruce grouse. Another species, Eim- eria bonasae was found in the cecum, rarely in the small intestine, of the ruffed grouse in Massachusetts, Labrador, Quebec and Alaska'". This form was also encountered in ptarmigan and later in sharp-tailed grouse. A third species, Eimeria dispersa, has been reported from the bobwhite quail, ruffed grouse, domestic fowl, turkey, pheasant and sharptail. The species encountered in New York are probably the Eimeria bonasae and Eimeria dispersa. Description. Coccidia are microscopic protozoan parasites which, in birds, infect the lin- ing cells of the intestine. The mature forms of this parasite, the oocysts, are spherical or oval cysts with distinct cell walls. Coccidia are distinguished from each other by the size and shape of the oocysts and the number of included sporozoites. Eimeria angusta is an elongate form measuring from 16.5 to 17.5 microns^ in width and 27 to 33 microns in length. E. bonasae, almost spherical in shape, measures 21 microns in diameter and is found in the caeca and rarely in the small intestine. The oocysts of E. dispersa are about 19.8 x 17.7 microns. Life Cycle and Dissemination. Detailed accounts of the life cycle of coccidia are available elsewhere". Very briefly, the hfe cycle may be outlined as follows. The microscopic oocyst, when discharged with the feces, undergoes internal growth. There develops, within a period of from 24 to 72 hours, four spores, each of which in turn divides into two elongate sporozo- ites. When the oocysts containing the sporozoites are eaten by a bird, the cyst wall is dissolved by digestive enzymes and the sproozoites are liberated. They inunediately proceed to penetrate tile lining cells of the intestine. Ntiw begins a sequence of events characterized chiefly by the growth of the parasites and their tremendous multiplication at the expense of the intestinal Un- ing cells of the host. The cycle of growth terminates in the (le\olii]nneiit of another crop of oocysts. Most of the work on avian coccidiosis has been carried out with domestic chickens. In gen- eral the following factors with regard to the spread of coccidiosis would apply to all species of birds: * Se« ditruMJon of Diipair in Hanil-rt-arrd Griiuflr, p. 438. A A microo it • uoil of microicopic meaiuremenl which «quaU about 1/25,000 of an inch. PARASITES OF WILD GROUSE 429 1. Coccidia infecting one species of host seldom infect another. E. dispersa is an exception to this principle since it has been reported from a number of birds. 2. Oocysts under suitable conditions may survive for a year or more on the ground. 3. In all likelihood, adult birds previously infected may serve as carriers of infection for susceptible birds. 4. In some types of infection, there is an age resistance exhibited by the host. 5. The severity of infection is determined, among other things, by the number and species of the infective stages eaten. 6. Recovery from infection results in a state of immunity in the host. The extent and nature of this immunity and the promptness with which it is brought about depends upon the species of coccidia and the severity of the initial infection. There is no reason to doubt that these factors operate in the spread of coccidiosis in ruffed grouse. Obviously, until detailed laboratory experimentation and further field observ atiuiis are done with coccidia of grouse, one may only conjecture on the exact operation of these factors. Pathogenicity. No instance of severe coccidial infection has been encountered by the Investigation. For the most part, the presence of coccidia has been detected by the examina- tion of smears from tlie intestinal mucosa. No attempt has been made to identify the species of oocysts encountered. It is interesting to record that on one occasion during the early attempts to propagate grouse in confinement, fecal examinations* carried out as a part of the routine examination, revealed large numbers of oocysts in the feces. Despite this, the half- grown birds showed no symptoms of disease nor was there any clinical evidence that intestinal inflammation was present. Incidence and Importance. As is evident from table 64 and table 67, the greatest incidence of coccidia was found in grouse chicks less than three months of age. In this group 10.1 per cent of the chicks were found to be affected. On the other hand only 0.7 per cent of the birds over three months were found infected. A further breakdown of the data indicates that during the first two months, the rate of infection remains alxiul the same, 12.2 per cent and 11.9 per cent. The incidence of infection in the birds from two months and older takes a sudden drop to 1.4 per cent and decreases still further until no ex idence of infection was found in the fall and winter months. It would seem from tiiis data, that probably a pmnipt infection of susceptible chicks occurs early in life and that an immunity is quickly built up so that subsequent exposure results in little, if any, infection. Tyzzer"' suggested this during his early work on coccidiosis in game birds. The fact that very young chicks feed largely on food on the ground probably accounts for considerable ingestion of oocysts. As the feeding habits of the birds change and less pick- ing is done from the ground, fewer chances for infection occur. The relatively low incidence of coccidia and the fact that pathological conditions have not been observed in ruffed grouse as a result of coccidial infections, make it obligatory to exclude these parasites from serious consideration as a factor in grouse mortality. Though early grouse breeders" reported coccidiosis to be one of the serious problems en- countered in artificial propagation of recent years, the number of cases observed in captive grouse raised at the Research Center has been almost negligible. * Sugar flotation technique. 430 PARASITISM AND DISEASE L\ RUFFED GROUSE Blood Parasites The parasites that have been identified from the blood stream of New York grouse are of only one type. Microfilaria. From Ontario, Clarke '■■ '• "' has reported two malaria-like para- sites, Leucocytozoon and Haemoproteus and a spindle-shaped protozoon, Trypanosoma. The last two have also been identified from grouse in Michigan. Leucocytozoon attacks young grouse with serious consequences, according to reports. Microfilaria Distribution. These blood parasites have been found in grouse in Ontario and New York. Since specific identification could not be made and no adults were found, it is impossible to say whether the same worms exist in other birds. Description. Microfilaria are microscopic roundworms which occur in the blood. They are presumably the immature stages of tissue dwelling roundworms. These forms have been found in the blood smears of many birds, in which no adult filarial worms have been found. Life Cycle and Dissemination. The group of parasites to which the microfilaria belong de- posit living embryos in the blood streams of their hosts. These are taken up 1)\ blood-suck- ing insects when they feed and are transmitted to other hosts by their bites. Patkogenicity. So far as is known these parasites do not harm their hosts. Incidence and Importance. Microfilaria were not found in chicks and occurred in only 1.2 per cent of the adult. There is no reason to believe they are of any importance to grouse. External Parasites In the wild, the external parasites most consistently found on grouse are the ticks and louse flies, although lice and mites are sometimes encountered. As has been noted in table 64, ticks have been found on about 12 per cent of the birds during the summer months while louse flies occur on from 2 to 7 per cent, depending on the age of the birds. It is only during the summer months, when special collectors are in the field, that the freshly shot grouse are placed in wax paper bags immediately on collection. The incidences given above are based on these summer collections, although birds received throughout the year are examined for these parasites and spring and fall occurrences have been recorded. Ticks ( Haemaphysalis chordeilis and H. le peris- paluslris) Distribution. As has been indicated at another point in this chapter, the majority of the ticks identified from grouse during this Investigation, have been bird ticks (Haemaphysalis chordeilis). Some rabbit ticks (II. le ports- palustrisj have been taken and probably some num- bers of nymphal stages of ticks, which could not be identified to species, belong to the latter group. The bird tick is also resident on other birds across the country, including chickens, turkeys, spruce grouse and songbirds. Il has been implicated in the transmission of tularemia among sage hens. The rabbit tick is found on cottontails, varying hares and jackrabbits as well as on a number of other species of mammals and birds throughout most of North .America. Ticks were found on grouse in all three regions in New York but they were more abundant in the Adirondacks than in other parts of the State. Description. The licks found on grouse manifest different si/cs and shapes depending on their age and state of engorgement. As they suck blood from their hosts they increase in size, the largest being about % of an inch long and roughly spherical or ovoid. The females are PARASITES OF WILD GROUSE 431 larger than males and the color when engorged is a reddish brown. Young and unengorged individuals are much smaller and lighter in color. Identification of ticks to species requires the services of specialists in this line. Life History and Dissemination. From the eggs of the ticks which are deposited on the ground, small six-legged "seed" ticks hatch. These climb upon vegetation and wait for contact with a suitable host. After attacking themselves thereto and feeding for some time, these larvae drop to the ground and undergo a period of relative inactivity. From this they emerge as 8-legged forms known as nymphs. These forms also must contact a host on which to feed. Later they too, fall to the ground where a final transformation into adults takes place. The adults become attached to a third host, on which mating takes place. Pregnant females like- wise return to the ground to lay their eggs after which they shortly die. Although adult ticks have their preferred hosts, they are sometimes found on a number of species which they do not inhabit commonly. In general, the larvae and nymphs are not as particular as are the adults in the selection of hosts. They may therefore be found on many different animals. This ability to utilize a number of different species is of particular importance in the maintenance and dissemination of ticks which do not remain on the same animal through all stages, since in dropping to the ground to moult the parasites run a risk of not being able to contact an- other host when the time comes for subsequent engorgement. Pathogenicity. Ticks gain their sustenance by sucking blood and during this process may set up considerable inflammation and irritation at the site of their attachment. They are pre- sent on both chick and adult grouse during the summer. BIRD TICKS ( Haemaphysalis chordeilis) ATTACHED to head and neck of adult grouse 432 PARASITISM AND DISEASE IN RVFFED GROUSE In only one case, however, was the death of a l)ii(J attiil)uled to these parasites. On June 6, 1940, an adult female grouse, somewhat emaciated, was found near Glens Falls in a dying condition. It was sent to the Research Center and died enroute. Aine adult bird ticks, all fully engorged, were found attached about the neck close to the head. It was apparent that their presence had caused much irritation. In fact, many feathers had heen lost from the region. Complete examination of the bird revealed conditions indicative of starvation. No food or grit had been taken in some time, the gizzard was flabby and death was attributed to malnutrition developed as a result of this external parasitism. The relation of ticks to tularemia in birds and mammals has been considered elsewhere*. Incidence and hnpurtance. Only during the summer has accurate data been obtained to indicate the incidence of infestation and the number of ticks per bird. Grouse in the Adiron- dacks were more frequently infested tlian those in the other regions of the State and each in- fested bird in that region harbored more ticks than were observed on individual hosts in other regions. The heaviest infestations were observed and the greatest numbers of grouse were found to have ticks during July. These parasites were found in about the same degree each year. Xo cases were observed in which the number of ticks exceeded 300, though much hea- vier infestations have been reported in the mid-west. The first record of ticks obtained by the Investigation was that of two from a captured chick, 12-14 days old, during the summer of 1930 and were identified as //. chordeilis. The earliest date on which ticks were found on adult grouse during the Investigation was May 4th. Though no accurate count after the end of August is available, field observations seem to indicate that these parasites are rarely found on grouse as late as the hunting season in Octo- ber. Adult ticks are readily observed on account of their size. Many amateur observers have therefore considered them to be important causes of grouse decimation. Yet, evidence of ex- tensive damage has not appeared during the course of the Investigation. This is another case in which the obvious parasites prove to be less important than the inconspicuous ones. Louse Fly (Lynrhia americana) Dislribution. This parasite commonly found on grouse during the warmer months, is also frequently encountered in New York on great horned owls and on marsh, red-tailed, and broad-winged hawks. In other localities in North America, a number of other hawks and owls have also been reported as hosts of this fly. Description. Though these insects have been called "flying ticks" the\ are in reality, flies which are adapted to an ectoparasitic existence. They have flattened brownish bodies and long wings which bring their t(jtal length to almost ^^ i'K'h. Since they quickly lea\e the body of a dying bird, they are seldom observed uidess the freshly shot gnmse is immediately located and retrieved, lender these conditions, they may be noticed h\ the sportsman during the hunting season. Lije Cycle and Dissemination. Little is known of the life cycle of tliesc insects. In flies of this group (Pupii)ara) the eggs are not laid by the female but are retained within her body where they hatch and develop to the state of pupation before they are deposited. It has been suggested that the adults do not survive the winter in temperate regions. If this is the case, reinfestation of non-migratory birds must take place each spring when the migratory hosts return. Certain is it that they have not been encountered in the spring earlier than March 26. Pathogenicity. No deleterious effects have been observed as a result of iMfcstatiim with these * See dilcusBtun of Relation of DiscaBCB uf Wild Mammal* lu Grouie, p. 416, PARASITES OF WILD GROUSE 433 external parasites. Related flies (Lynchia hirsutaj and ( Lynchia fusca) have been shown to be vectors of malaria-like diseases of CaUfornia valley quail^-"". Another form, Pseudolynchia canariensis, serves as the intermediate host of a related blood parasite of pigeons^'. Al- though Clarke' has reported a Haemoproleus infection in grouse and suggested the louse fly as the possible vector, no conclusive evidence exists to confirm this view. Incidence and Importance. The low incidence of this parasite and the little trouble which it apparently causes, seem to indicate that this organism is of no great importance in limit- ing grouse abundance. Parasites Not Encountered During the Investigation During the course of investigations on grouse carried on in other parts of the country, a number of parasites have been recorded which have not been encountered in Aew York or which appeared so infrequently as to make evaluation of their role impossible. Important among these is a parasite of the red-blood cells of grouse, (Leucocytozoon bon- asae), first noted in Ontario and since found in Michigan and Maine. It is presumably trans- mitted by the bite of the black-fly (Simulium venustum). Clarke' in studying Canadian grouse, considered it "significantly associated with the cyclic diminution" in numbers. O'Roke, who has made a study of the malaria-like disease of ducks^ caused by Leucocytozoon anatis, also regarded this grouse parasite to be pathogenic in young birds^"', though he did not study it during a period when die-off was apparent. During the Investigation, blood smears from 781 grouse were examined for the presence of this parasite. Of these, 214 were not taken during the summer months and therefore might not be expected to show the parasite. Of the smears taken during the summer, 106 were from adult birds and 461 were from chicks. None of the smears examined revealed the presence of the parasite. In addition to the blood smears, livers, in which certain stages of Leucocytozoon are found, were saved for miscroscopic study. Of the 195 livers examined, 117 were from chicks and 78 were from birds over three months old collected in October and November. These likewise were negative. The blood smears and livers from birds taken in many parts of the State, though the majority of the specimens came from Essex and Lewis Counties in the Adiron- dacks, Tompkins County in the Southern Tier and from southern Albany County at the northern edge of the Catskills. A related one-celled animal (Haemoproleus sp.) which inhabits the red blood cells of ruffed grouse was found in Ontario by Clarke" in the summer of 1935. Nothing is known of its effect on grouse. A parasite of the same genus is transmitted to California valley quail by louse flies^ and it has been suggested that the same process may occur in ruffed grouse. Still another protozoan (Trypanosoma) which lives in the fluid portion of the blood but not in the blood cells, has been reported from ruffed grouse in Michigan'^' and Ontario'' but no disease condition is attributed to its presence. Certain one-celled intestinal parasites have been described from New England grouse, though they apparently are associated with no conditions harmful to the bird. These organisms move by means of whip-like structures and are commonly known as flagellate* (Trichomonas bonasae)^'", (Cyathosoma striatum), ( Ptychostoma bonasae )''' . Shillinger and Morley''' have reported that similar organisms, resembling Trichomonas gallinarum, have been observed to cause a highly fatal disease of young quail and grouse. 434 PARASITISM AND DISEASE /A RUFFED GROUSE Several species of flatworms known as flukes, have been reported from scattered localities throughout the range of the grouse. One ( Hannostomiim pellucidum) was originally found in the cecum of a robin'"" but has been encountered in ruffed grouse in Minnesota"*, New York, New Hampshire"', and possibly Ontario'. Another very minute worm ( Leucochloridium pricei) first found in the rectum of an Alaska spruce grouse"^" was recently found in ruffed grouse in New Hampshire"'. Neither of the above parasites appear to damage the birds. Prob- ably both species are obtained by eating certain land snails. Other intestinal flukes reported for ruffed grouse are Glaphyrostomum from Labrador"^ and a spiny-mouthed form i Echinoparyphium aconiatum ) from Minnesota""". The latter was first described from tlie lapwing'"" in Europe and is probably accidental in grouse. Some of this group of flatworms occur in other organs. Ly perosomum monenteron, which was described from the livers of robins, bluebirds and kingbirds"", has been found in grouse in Minnesota. In the muscles of grouse in that State, immature flukes", Agamodistomum sp., have been en- countered. Proslhogonimus macrorchis, the oviduct fluke of poultry and pheasants, has been reported from the bursa Fabricii of grouse in New Hampshire""^'. A tapeworm (Davainea proglottina) which is generally considered to be a parasite of chickens and turkeys has been found in ruffed grouse in Labrador"' and New Hampshire"'. The fact that no poultry existed in the former region where this worm occurs poses an in- teresting question. Is it typical of chickens and transmissible to grouse or is the wild bird the normal host? Two other poultry tapeworms have also been reported from grouse (Choanotaenia injuiidi- buluin and Hy/nenolepis carioca). These were found in Minnesota" and New Hampshire", respectively, the identification of the latter being tentative. Davainea telraoensis, a tape- worm known also in the capercailzie in Switzerland, has been encountered in ruffed grouse in Michigan'"' and Ontario". In the mid-west""' "' a roundworm parasite (Oxyspirura petrowi) occurs under the nictitating membrane of the eye of grouse occasionally causing considerable inflammation. An intestinal roundworm (Contracaecum) has been listed** as a parasite of grouse but is probably an acci- dental occurrence because worms of this genus are generally found in fish-eating birds and mammals. The strawberry worm (Tetrameres americana) whose common name is suggested by the almost spherical, blood-red form of the female, is found in the glandular stomach of chickens and l)obwhite quail and has been experimentally transmitted to grouse, pigeon and domestic duck by feeding certain species of infected grasshoppers'"'. There is nothing to indicate that it is a natural grouse parasite or that it is of importance in the wild. Nema- tode cysts, (Pliysaloplcra) found in the muscles of grouse in Minnesota''' are undoubtedly accidental and innocuous. Rodent mites ( Laelaptinae) have liccn rc|)(iiled from grouse in New Hainjishire"'" and harvest mites (Trombicula microti) from Ontario'" while in both ihcse localities feather mites have also been found'"' *". In the summer of 19,32. a female grouse was lhi>hi'(l from her nest in Otsego County, New York, revealing the presence of hundreds of miles wilhin llic nest and covering the eggs. Un- fortunately there were no facilities for their eollei linn ;iiul lliey were, therefore, not made available for identification. The parasites of this group, however, are thought to cause but little if anv trouble among grouse. The northern fowl mile, (Liponyssus sylviaruni ) on the other hand, has been reported to DISEASE IN HAND-REARED GROUSE 435 have caused the death of captive birds in Maine'" while the scaly-leg mite fCnemidocoptes mutans) which burrows under the scales of the legs, may cause severe inflammation and lameness. Both species are ordinarily poultry parasites. The latter also occurs in pheas- ants and was recently reported from ruffed grouse in British Columbia". Among the insect parasites occurring occasionally on grouse are bird fleas (Ceratophyllus diffinis)^" and bird lice (Gallilipeurus cameratus", Lagopoecus perplexus^, Goniocotes sp/"' and Menopon sp.). Although the above list may seem long and formidable, in actuality grouse probably do not harbor more parasites than do most other game birds. As has been pointed out, in most cases, large numbers of any given species do not appear in individual birds. It is sel- dom that one bird harbors more than a half dozen difi^erent parasitic forms. It is also im- portant to recall how few of the organisms mentioned during preceding sections have been demonstrated to have disease producing proclivities. The fear that the numbers of grouse in the coverts are seriously affected by their internal and external inhabitants has not yet been substantiated. DISEASE IN HAND-REARED GROUSE Early experience in the hand-rearing of grouse dcnioiistrated that discouraging losses re- sulted from disease". The maladies observed most comnKinly were blackhead (enterohepa- titis), stomach worm infection, ulcerative enteritis ("quail disease") and coccidiosis. The presence of these afflictions was probably due partly to the fact that the birds were raised on the ground or in some form of lifter rather than on wire, partiv that they were usually in contact with ])oultry, directly or indirectly. In some cases, indirect connection with other game birds, particularly quail, may have been responsible. Knowledge concerning the means of transmission and spread of parasites and disease has grown. Practical methods of prevention and control based on this knowledge, have been adopted. The use of hens as foster mothers for young grouse has been abandoned. Rearing of birds on the ground is now considered to be not only unnecessary but dangerous. These steps have led from a disheartening period of pioneering to the present phase in artificial propagation when infectious disease is less of a problem than certain aspects of incubation, brooding and nutrition. An understanding of the methods used in hand-rearing grouse at the Research Center as described in Chapter XI, Artificial Propagation of Grouse, should logicallv precede the con- sideration of the disease conditions encountered in captive grouse. Described in a nutshell — the eggs are artificially incubated and the newly hatched chicks are transferred to battery brooders with wire floors. At the end of ten days they are placed on wire in a colony house where they remain until they are 6 weeks old, at which time they are transferred to outdoor pens again equipped with wire floors. The nearest poultry are one-half mile away and the only other birds kept at the Research Center are waterfowl and pheasants. The sanitary precautions which are an inherent part of the propagation program elimi- nate worm parasitism entirely and reduce even highly contagious diseases, such as ulcera- tive enteritis and blackhead, to a few cases a year, while tuberculosis, aspergillosis and coc- cidiosis are so rare as to hardly require consideration. The use of wire-bottomed pens to prevent contamination of food and water with droppings is the procedure to which this con- trol is largely attributable. 436 PARASITISM AND DISEASE IN RUFFED GROUSE It has been pointed out that, during recent years, none of the pathological conditions en- countered in the wild* have occurred in grouse hand-reared at the Research Center, and none of the diseases which occasionally appear in captive grouse* have been found in wild birds. It has been observed that the infestations with bird lice, when they do occur, are heavier in the captive birds than in the wild ones. The lice involved, however, are not the same species. The ones which occur on the hand-reared birds have pheasants as their ordi- nary hosts. The heavier infestations are undoubtedly due to the lack of dusting facilities in the cages. In the following sections, the important specific diseases of ruffed grouse in captivitv are discussed. Three of these, "quail disease", blackhead and tuberculosis, make their primary invasions in the digestive tract, though other organs may later be affected. Two, aspergillosis and air-sac mites, are typically disorders of the respiratory tract and its appendages. One is a nutritional disorder which affects the leg joints. It should be indicated that certain other pathological conditions appear in hand-reared grouse, though no specific disease-producing organism is held to be responsible for their oc- currence. For instance, congestion of the lungs or pneumonia, which is observed in young chicks, generally follows exposure to over-heating or chilling. This may result from inade- quate brooding facilities. There is also a condition in which urates, substances ordinarily eliminated by the urinary system, are retained within the body and precipitate in the kid- neys, liver and heart. Since this disorder can be produced experimentally by altering the diet or environment, it is difficult to identify its cause. In the following sections, detailed descriptions of symptoms of the various diseases have been omitted because of their undependability in the diagnosis of disease. Sick birds usu- ally tend to present the same appearance and behavior when afflicted with any of a variety of disorders. Positive identification of pathological conditions is best made by post-mortem ex- amination of birds which died of disease or birds selected from pens where infection is sus- pected. The several diseases which afflict hand-reared grouse may now be taken under specific consideration. "Quail Disease" or Ulcerative enteritis Cause. The causative agent of this infection is still in doubt. Morley and Wetmore*" isolated an organism which they believed to be the cause and named it Corynebacteriiim per- dicium. Recently Bass'" reported that the true causative agent was an anaerobic organism. Neither of these claims have been confirmed b\ nuraiit and noil"" wlm \mmc iiiuiblc to deter- mine the cause of this disease. Nature of the Disease. The malady may be described as an acute contagious disease tliai allacks the ceca and intestine of quail, grouse, chukars and turkeys. Outbreaks in the two former species have occurred only in artificially propagated birds although Green and ShilliiiL'cr'"" in Minnesota have re|)orted finding ulcerative Ul' A.SI'EIU.ILI.U.SLS I.N LI .N(. AM) K1I)\K\ OF AIH LT OHOLSE DISEASE IN HAND-REARED GROUSE 441 When the disease occurs in young pouhry, during the brooding period, it has an acute course for death commonly occurs within a few days after exposure. This rapidly fatal form of the malady is extremely rare in grouse. Most cases which have been observed at the Center have been chronic and confined to adult birds. Aspergillosis is a decidedly infrequent finding in grouse autopsies. Dissemination. Dark and damp conditions either in the bird's quarters or in places of grain storage are conducive to the development and maintenance of the mold. Control. Procedures which foster dry, clean, uncrowded housing of birds and feed should be employed. Air Sac Mites i Cytoleichus nudus) Cause. The air sac mites are very small spider-like animals, found in the air sacs and less often in the bronchioles, lungs, trachea and even in the hollow bones of grouse and other game birds, chickens and |)igeons. The parasites appear as tiny yellowish-white spots. The males measure 450 x 300 microns and the females 300-600 x 400 microns. Nature of the Disease. Light infections of these mites produced no gross pathologic changes. If the birds had just been killed or had just died, the tiny mites could be detected by their slow movement over the air sac membranes. In heavier infections the membranes became much thicker than normal and leiided to become nicire opaque. In a few severe cases, pneumonic areas in the lungs with catarrhal exudate in the bidtichi and lower trachea were found. Large numbers of eggs and adult mites were seen in wcl prc-paraliotis of mucus takcti from these places. Very few hand-reared grouse succumbed In tliis infection. Dissemination. Little is known about its life c\(le or natural method of transmission from one host to the next. The females lay eggs which contain larval mites ready to hatch. In the cases encoutitt'rcd in artificiallv reared grouse, infection was believed to have been acquired from chickens which, in the early days of the Investigation, were used for hatching grouse eggs. Control. Discontinuance of the use of chickens as foster mothers and prompt isolation of all grouse that appeared dumpy served to eliminate this infection. Slipped Tendon. Hock Disease or Perosis Cause. Work by Wilgus and co-workers™' in 19.S7 and confirmed since then by many others, indicates that the basic cause for this nutritional disease is a lack of manganese in the ration. Nature of the Disease. This disease, fairly common in domestic poultry, is characterized by the shifting either to one side or the other of the large Achilles tendon as it passes over the hock joint. There resuhs a curving of the shanks and a twisting of the leg out of the normal position. The birds not only become lame but in many instances are obliged to hobble along with the hock dragging along the ground. Only a small number of grouse chicks succumbed to this condition although it always seemed that the largest and fastest growing chicks were most likely to come down with perosis. The disease has not been observed among adults. Control. Prevention of this condition is effective when sufficient manganese is present in the ration. Domestic poultry require 50 parts of manganese per million. 442 PARASITISM AND DISEASE /A' RUFFED GROUSE THE ROLE OF DISEASE It can be seen from the preceding pages that the relation of parasitism to disease and disease to the life of the ruffed grouse is difficuU to evaluate. Many of the parasites do not produce disease and those which do are not widely distributed throughout the range of the bird. Though it has long been asserted as a generality that the limits of game populations are probably set by disease^", clear cut examples of this are not easy to find. It seems logical to assume that transmission from animal to animal takes place more readily as the population increases. Furthermore, even though the increase of virulence of certain disease producing organisms by rapid passage through susceptible hosts is demonstrable, the actual occurrence of these phenomena in nature is not often observed. Under conditions obtaining in human populations and in concentrations of domestic animals, these processes may be followed, but to explain density fluctuations in wildlife on this basis, without more definite information than that available to date, seems inadvisable. Whether disease reaches dangerous levels in natural grouse coverts is a matter for specula- tion. The question is by no means solved by pointing to the absence of other factors to which "die ofFs" can be attributed. The difficulties encountered in studying the role of disease in wild animals foster the temptation to make broad generalizations the basis for interpreting prob- lems which must be approached from a factual standpoint if sound conclusions are to be drawn. The possibility that local variations in abundance may occur and the fact that complete in- formation on relative abundance cannot be obtained for all of the areas from which birds are collected, further handicaps interpretation of autopsy findings in terms of the relation of pathological conditions to the grouse population An additional matter for conjecture is the suggestion that, since, during periods of abund- ance, transmission of disease is probably facilitated, increased hunting pressure during such times would be of benefit in reducing the population to a "safe" level. As a corollarj' to this is the attitude that certain of the birds shot in fall might well die of disease or other causes during the winter, if they were not so harvested. Proper evaluation of these questions should also be based on sound observations on the actual rate and mode of increase of disease during increases in abundance and on the relation of the haphazard sample taken in fall hunting to the normal and diseased sectors of the bird population. The fact that disease does occur and on occasion accounts for deaths of both wild and captive grouse, necessitates its recognition as one of the factors limiting abundance. In this sense, it ran no more be removed from consideration than can weather, predators and the other items which in one wav or another influence the lives of grouse individuals and groups. Observation on pathological conditions during a period of over ten years, however, has revealed no disease of epizootic proportions which would account for a sudden and wide- spread disappearance of ruffed grouse. CHAPTER XI ARTIFICIAL PROPAGATION By Gardiner Bump ORIENTATION Background and Difficulties — Methods of Raising Grouse — Progress to Date — Results THE STARTING POINT Eggs from Wild Nests — Collecting and Transporting Grouse Eggs — Wild-Trapped Birds THE NATURAL COVER METHOD OF RAISING GROUSE Penning Birds and Providing Shelter — Feeding — Protection from Predators — Danger from Disease — Vitality THE INCUBATOR-BROODER METHOD OF RAISING GROUSE Caring for the Breeding Stock — Overwintering Grouse Breeders — pens and penning — foods and feeding techniques — handling the spring shuffle — weight and physical well-being — The Spring Period — selection of breeding stock — breeding pens and equipment — pen- ning and care of breeders — feeding breeding birds — mating and egg fertility — egg pro- duction and collection — controlling disease — The Summer and Fall Period — moulting and recuperation — The Incubation of Grouse Eggs — Grouse Mothers — Foster Mothers — construction of hatching coop and nest — care of eggs — -The Incubator Method — location of the incubator — type and care — incubator operation — measuring incubation progress — setting and care of eggs — hatching and transfer of chicks — Brooding of the Chicks — Brooders and Brooder Houses — Brooder Operation — Care of the Chicks — Feed and Feed- ing Techniques — Feather Picking and Cannibalism — Reducing the Danger from Disease and Predation — Rearing of the Young Birds — Rearing Fields and Units — Pens and Pen- ning— Feed and Feeding Techniques — Disease Prevention and Control — Care During Rearing — The Period of Social Adjustment. PRACTICABILITY OF ARTIFICIAL GROUSE PRODUCTION Production Equations — Factors Limiting Production — Biological Limitations — Scarcity of Obtainable Breeding Stock or Eggs — Lack of Trained Manpower — Cost SIGNIFICANCE OF ARTIFICIAL PROPAGATION 444 ARTIFICIAL PROPAGATION OF GROUSE GROUSE LIBERATIONS Transportation for Liberation — Method of Liberation — Marking for Identifica- tion— Vi'HAT Determines Slrvival — Source and Experience — Age — Physical Condition — Ability of Liberated Birds to Survive. <■ the means of sa\ing a sjiecies from extinction. (Jronse are still dillicult and costh to raise in capti\ itv 1>\ comparison with pheasants and (juail. For this reason the artificial production of grouse, eitlier to restock depleted i()\crts or as "birds for the gun" is at ]jresenl seUloni iu>lili('(l c\ii-pt where cost is not an impor- tant factor, where research ends are to be served, or where birds arc needed to reintroduce the species into a favorable habitat from which the\ have been extirj)ated. ORIE.NTATION Before proceeding with a detailed descri|ition of how to raise grouse in captivity it is well to consider brieflv the picture in general terms. \\ hat are the major problems one must face? What choice is there as to propagation methods'.'' What progress has been made to date in raising grouse in rapti\ily? Let us then get first a bird's-eye view of the situation. Background wn Difi-iculties One has but to turn to the first chapter of the book to see how clearly the game enthusiast of yesteryear visualized the productive potentialities that lay in artificial propagation. That they were seldom realized with grouse is due perhaps to the biology of the species rather than the men who tried tirelessly to raise these birds by adapting the methods that had proved so successful with pheasants and quail. 446 ARTIFICIAL PROPAGATION OF GROUSE Those who raised some grouse, and there have been many, found them to be tame and rather easy to manage. Though normally not a social species, at the New York Research Cen- ter they can be kept congenially a part of the year in flocks of up to 300 individuals. In cap- tivity, at least, grouse are polygamous. The eggs are not too difficult to hatch in incubators. Once the chicks have passed the month-old mark, they are as easy to raise as quail. But there are substantial difficulties to be overcome before mass production is economically feasible. For instance, physiologically, most female grouse do not seem to be gaited to lay the large numbers of eggs normally produced by the species which have been successfully raised in captivity. It is the occasional bird which lays more than 30 eggs that holds forth the promise of developing a high producing strain by selective breeding. The difficulty of synchronizing the mating period of the sexes and the tendency of the male to viciously dominate the female during much of the breeding period are frequent causes for trouble. An even more puzzling problem is presented by the chicks, many of which have a discouraging way of dying in the first month without any thus far detectable reason. Here again careful selection of the breeding stock to correct this difficulty may prove the solution. None of these troubles appear to be insurmountable. Eight years of selective breeding at the Research Center directed at increasing egg production, fertility, livability and social bal- ance, have shown some encouraging though many conflicting results. As every breeder knows this period is far too short to change even one characteristic when so many factors are involved. Quite frankly, a way of eliminating the early losses among young birds has not yet been found. That it may ultimately be accomplished is indicated by an occasional season during which 50 to 80 per cent of the birds hatched were raised to maturity. Some may be surprised that the much publicized susceptibility of grouse to disease is not included as a primary difficulty. In the light of today's experience, this factor is no more difficult to control in propagating grouse than in quail. In fact, unless intelligently handled, more birds may be lost, particularly in spring and fall, through the desire of some indi- vidual birds, usually males, to dominate their penmates. This tendency may be referred to as the "dominance complex". It may cause fighting and chasing, sometimes resulting in severe injuries to the subordinate birds, and it may also seriously impair the breeding potential of many of them. Unlike the primary difficulties listed above, these last two have been satis- factorily overcome. There has been one other substantial bar to progress, a characteristic not of the birds but of those who would raise them. Until recently, artificial propagation has been largely an empirical procedure. Here success depended upon the fortuitous blending of native intuition with a knowledge of the "secrets" of the trade. In many instances these have been well guarded. Although, when i)r(>p('rly ecjuippcd. it is ti>d;i\ no trick to raise these birds from eggs col- lected in the wild, little progress is likely to result from so doing, for every conceivable angle of this procedure has been thoroughly explored. It is lime indeed for the enthusiastic grouse 'HlM*.V»4AUM«fc«3^UjJ«i»*>4jSjl ■\-^Ajc2.5_..«v. >^x&•;UJ•.'4» ORIENTATION 447 experimenter to realize that the problem may not be solved by adopting a new type of hover or by adding v^rintergreen to the diet. In fact, little useful purpose beyond encour- aging public hopes and interest is likely to be served by continuing such rule-of-thumb experiments. Large-scale production of grouse in captivity apparently depends upon the ability of grouse breeders to produce, by selective breeding, a strain of birds which will lay a large number of fertile eggs which will, when hatched, produce chicks that can be reared to maturity with- out undue effort or loss. Perhaps a hundred individuals have raised a few grouse in cap- tivity, but each has been unable to translate early successes on a small scale into continuous large-scale production. Methods of Raising Grouse " There are but two main systems of raising grouse in captivity, the natural cover and the incubation brooder methods. Under the first, the birds are maintained on the ground in large covered pens under semi-natural conditions. In contrast with this, birds that are hatched in incubators, reared and held on wire and closely penned are said to be handled by the incu- bator-brooder method. Many combinations of these have been tried. The merits and drawbacks of each method are described later in some detail by way of presenting the whole picture, although the factor of disease has made the raising of grouse on the ground at best a precarious proposition. Progress to Date This phase of the Investigation was largely an outgrowth of the pioneering experiments of Dr. A. A. Allen at Cornell", although Hodge, Merrill, and Torrey had demonstrated pre- viously that grouse could be raised from wild eggs*. However, little progress had been made in overcoming many of the difficulties standing in the way of mass production. While trying modifications of the old and accepted methods for ten years, Allen realized that his first limitation lay in the susceptibility of grouse to certain diseases. He then made two pivotal advances. First he abandoned disease-carrying hens as foster mothers in favor of the then none too reliable artificial hovers. Secondly he developed a method of raising and wintering grouse on wire. * See itory of early grouse rearing experiments in Chapter I. 448 IKTIFICI 11. l'li()l'AGATI(>\ OF (;n<)LSE The comijaralive freedom fnim disi-asc thus gained provided the first real opporlunitv for finding practical methods of raising the chicks, maintaining thoni as hrcedors and stncKing the pnihlem of securing a large nuniher of fertile eggs. The prolileni \\as allac kcd on two fronts. 1 lie Iri\estigation established in 1931 a small field station in the western tiatskills near the \ illage of Har\ard. There, as a result of the confusion of methods emploxed by those then raising grouse, main of the earlier tech- niques were rec lieeked. The major effort was concentrated, howexer. on de\elo|)ing and testing new ideas for rearing grouse on wire in small artificially heated brooders. Coincidental with the work at tliis station, funds and equipment were provided Dr. Allen by the Investigation so that lie might continue his ouii Icuig-tiiiie and promising experi- ments at Ithaca*. liitrilitirr Hump MKI.l) STATION IN THE W KSTKHN (Alskll.l, AKil M \I>s W IIIIIK 1 III, IW I >l li, \ i ION's FIRST TESTS OK REAKING GROUSE UNDER SEMI-NATUR.\L CONDITIONS. AS WELL AS IN HKOODERS. WERE CMUUEI) OUT By 19.33 interest in scientific game |irii(lncti:)n had increased In ihe pninl uliere an I'.xperi- niental Game Farm was estaiilished at Dclmar not far from Albain. The grouse raising |)roject. as one of the Deparliiieiit s major experimental acti\ities. accordingly was trans- ferred to llie new station. l-]\ enlualK this unit lieeame tlie liiib of tile game research pro- gram of the ('.onser\ation Department until, in I9II). all activities were combined at the Farm |o form a lu'seanli ('enter. There grouse propagation became but one of the many projects. ihe details of tlie indi\l(hKd experiments carried on o\er the years are worth reading care- fidlv if one is interested in raising grouse. A sununarv of these has accordingly been included In the A|)pendix (p. cSTT). for space will peiniil here only a brief mention of the more produetixe projects and results. For two \ears at the Catskill Station, some liird> were iiKiiiilained on the ground. This amply confirmed Dr. Allen's conclnsion that the lii>t |)rolilem to (dri(|ner was to learn how to control disease. A real start had been made b\ rearini; and carr\ ing grouse on wire. * Scr Annual Rcimrlii »i( iNfw Ytirk .Sinle Cunitrr\ntiun Dt- |uirlinni4 ol tl»' .North American C.nmr C«»n(rrcnco l..r lOI.'U I ORIENTATION 449 The slogan then coined, "Never let their feet touch the ground, " proved valid at least to the extent of preventing epizootics of blackhead among old and young alike. Real strides were made in learning how to render impotent other limiting factors. The years from 1931 through 1937 were largely spent in learning how to raise and maintain healthy grouse on commercially obtainable feeds and in testing out a host of jnomising incu- bation, brooding, breeding and disease prevention practices. The artificial incubation of grouse eggs presented few serious problems although exten- sive tests both at the Research Center and at Cornell"'' were conducted to determine the proper technique. Precedent was shattered by substituting for the tinic-hDiiorcd chick did of hard-boiled eggs, clabber, and fly larvae or ant egg, an easily purchased game bird feed to which was added white fishmeal and milk, liver and brewers yeast all carefully dried. Subsequentlv. equally satisfactory results were secured by utilizing a connncrcial turkey mash supplemented by grains and green food. Brooding, too, was much simplified. The small hovers and wire runs used in 1931 were soon superseded by a large, all-purpose pen with an enclosed eleclricalh heated hover in one cor- ner. Since this was not altogether satisfactory, it was soon abandoned in favor of colony brooder houses. Several types and sizes were tested. As starting units, baltcrv brooders ada])ted for grouse were develo|)cd to facilitate raising the birds during the first 12 days following hatching. The inventive genius of hover manufacturers was then at its In-sl and dav-old grouse chicks were carefully exposed to all commoid\ used types, sizes and shapes of hovers with c<]ual impartiality. A low, rectangular box-type "still air" hover proved most adaptable though the difference between it and others was not great. Birds were brooded in groups of from six to o\er IIM) uilli ImsI n>iill> nlilaiiicd from small groups of not t)ver 25. It has alwavs been common practice to shift the young birds at about six weeks from brooders to rearing pens. Of tiicsc. many sizes and designs were tried out and the mini- mum requirements for rearing and caring for vnung birds, while thus confined, wi're determined. The opinion that adult grouse should be overwintered, full-winged in large pens under natural lover conditions, has liecii held by manv who have attempted to rear this bird. This method was given an extensive trial over a two-year period. Some trouble \\\\.h disease was experienced but losses from predation. accident and escape were prohibitixc. Nor were more eggs produced bv this group than h\ their close-penned companions. Other pens, including those of the all-purpose t\pe. so constructed as to facilitate holding the breeders in large groups on wire, were also gi\en thorough tests. Eventually an efficient overwintering pen. accommodating up to 300 breeders, was constructed. Because the birds could fly back and forth it was appropriately named the "flying pen." Throughout all the years much eflort was concentrated on the difficult problem of securing large numbers of fertile eggs. Breeders, breeding pens, combinations of breeding birds, food and care, all play a part in this problem. It was determined earlv that wild birds when reduced to captivitv were usuallv poor 450 ARTIFICIAL PROPAGATION OF GROUSE breeders by comparison with hand-raised grouse, although the former did lay some eggs while penned under semi-natural conditions at the Catskill Station. Even the number of fer- tile eggs laid by the birds raised in captivity was found to vary widely. In an effort to locate the reason for this, a study of penning conditions was undertaken. It was found that birds placed on the ground under semi-wild condition did no better than those on wire. The all-purpose pens previously mentioned proved quite acceptable to breeding birds. Much smaller pens were tried, but difficulties increased, resulting in a higher ratio of infer- tile eggs. To save steps, in 1937 a multiple-unit breeding pen was constructed, which, with few modifications, has proven quite satisfacto Looking towards the solution of the same problem, many combinations of breeding birds were tried out. Grouse are polygamous in captivity. Combinations of one male with up to five females in a pen have proven moderately satisfactory, although best results have been obtained by penning the birds in pairs early in March. Also, females were penned alone, fertile eggs being secured by introducing selected males which were in the mating cycle. Suc- cess proved, however, to depend largely upon the experience and capabilities of the attend- ant and so this latter practice, as a standard procedure was discontinued. After thoroughly testing several combinations of foods, a breeding mash and a grain mix- ture commercially produced for game birds were selected for feeding breeders. Begun in a small way in 1933, the planned selection of breeders for high egg produc- tion, fertility and for livabiUty in the resultant chicks was made a major project in 1936. Since then all three items have shown a slow and irregular but encouraging trend upwards. Disease has caused the termination of many a grouse experiment, though, contrary to gen- eral belief, grouse probably are no more susceptible to it than are bobwhite quail. It was learned early that to confine young grouse, even on fresh ground, was to invite trouble from disease. Some old as well as young birds died from blackhead or from ulcerative enteritis (quail disease) in the large covered pens at the Catskill Station even though great care was taken to avoid bringing in these ailments and no poultry had previously occupied the site. Both these diseases likewise caused considerable loss during the early experiments even when the grouse, old and young, were held on wire. Chemical preparations reputed to be disease-inhibiting when placed in food or water, were found to have little, if any, beneficial effect. Nor did "rubber glove" sanitation prove effec- tive in curbing epizootics of ulcerative enteritis and blackhead in the pens. Eventually, simple and satisfactory methods of prevention and control were worked out. Results To some who may, in a few minutes, read this summary of 12 years of experiment and glance at tables 179 to 183 in the Appendix, the story will not be complete without realizing that excellent brooding birds have been produced whose forebears, to the eleventh generation, have never touched their feet to earth. Nor have they been fed on aught save commercially prepared foods, interspersed with leaves of apple, dried alfalfa or needles of pine or hem- lock to furnish the requisite roughage. During the course of these experiments a total of l.WJ birds has been raised to maturity. Of this group 191 have been utilized in liberation experiments. From the rest, year after year, have been selected superior breeders from which must eventually come a strain of birds psychologically and physically adapted to withstand tho limitations and respond to the artificial stimulations of captivity. THE STARTING POINT 451 THE STARTING POINT Irrespective of whether natural cover or more artificial propagation methods are to be employed, the question most frequently asked by those who would breed grouse is, "How can a start be obtained?" Hand-raised stock is still scarcely obtainable at any price, thus limiting one to the securing of eggs from wild nests or from wild-trapped breeders. Eggs from Wild Nests Early experimentors, with few exceptions, started with the collection of a clutch or two of eggs from the wild. Nests were usually discovered by accident, for. unless one is skilled in the art, they are difficult to find. The Investigation, however, saved time by requesting the cooperation of game protectors, forest rangers and interested sportsmen. Advertisements in local papers inserted periodically for several years were responsible for the reporting of over 370 nests to the Investigation for purposes of study or of collecting the eggs. Difiiculties with adverse public opinion may be overcome if the experimental nature of the project is carefully publicized in the region from which the eggs are to be collected. Collecting and Transporting Grouse Eggs The nest location, when found, should be carefully marked to avoid subsequent confusion in refinding it though the immediate surroundings should not be disturbed lest predators thereby be attracted. It is not necessary to gather unincubated clutches for, if carefully handled, grouse eggs may be transported several hundred miles with no apparent ill effect even during the early stages of incubation when they are most sensitive to change. This may be completed with least likelihood of loss, however, just before hatching. Most of the eggs collected by the Investigation in the southern part of New York were transported between May 20-25. For several years great care was exercised in cidlpcting wild eggs and in transporting them overland or by plane over distances up to 200 miles. Upon collection they were placed in a pail half-filled with lukewarm grain and carried thus from nest site to a waiting car. There they were transferred to a well insulated five-gallon kettle, in the bottom of which heated grain had been placed. Over this were set a series of beaverboard trays in which the eggs were placed between layers of cotton batting. A thermometer was inserted in each layer of eggs. This was consulted as often as necessary to make certain that the tempera- ture did not rise above 100° F. or drop below 80° F. The container, open at the top except for a loose fitting beaverboard cover, was set on the car cushion or. if the road was unusu- ally rough, suspended by springs above the floor not far from the heater. By closing the car windows and turning on the heater, temperatures satisfactory to the eggs, if not to the driver, were maintained almost indefinitely. Later, heater and egg con- tainer were boxed in, thus permitting the car windows to be left open. At the height of egg collecting, several containers were sometimes delivered to a nearby airport from which they were flown to the Research Center by the Department's airplane. 452 ARTIFICI I/. PROP\GATIO\ OF GROISE Almiist iiiiifiirml\ iho eggs seemed to arrive in pxtelleiit ri»iidition. Eventualh . however, it was clis((i\crcd that, if thev were gathered within a period of three or four davs prior to hatcliing. no apparent harm resulted from transporting them in an unheated container carried on the front seat of the car. .At this late period incubation has apparently progressed to a |iiiinl where sufficient internal heating is generated to allow the eggs to resist cooling for IN THIS KKTTI.E, I\fa BATKD CROLSE EGGS, Bl RIED BETWEEN LAYERS OF COTTON, WERE CARRIED SUCCESSFULI.V OVER DISTANCES IP TO 2l)() MILES periods of six lip 12 iioins. In fart, one clutclL cold u|ioii collfctiori and tiioughl to he dead. w'as placed on .'^alurda\ noon in a refrigerator at a tcnipcraturc not far alio\e freezing. On the following Monda\ morning tiie clutch was removed for examination. The first egg to lie o|)cncd rexcalcd a lixc emlir\o. The remaining seven eggs were inuuedialelv jjlaced in an iniiihalor and >(\cral (hicks subsequenli\ were hatched. None li\ed for long, liow- e\cr. for ari\ nnusnai strain placed u|)on the cmlirMi usualh is rc(li'( ti'ii in lowered halcli- aliililj oi tlie egg and in a poorer livahiiity in tiie chick. W iLn-'ri!\i'ri:n Birds When (dtirined. w ild-trajjped hirds ma\ liecome lame in time, but are not dependable as egg producers. Most of them are imported from Canada, in this connection certain Fed- eral atwl .'^latc |ji'rmils coxciing liiipor lalion and |)Ossession aic iimkiHv icijiiircd. The birds should be secured as far in advance of the breeding season as possible and plaicfl in pens on wire in close proximit\ to man to facilitate the genlling-down process. The THE NATURAL COVER METHOD OF RAISING GROUSE 453 "all-purpose" pen hereafter described is quite adaptable to this purpose. To prevent injury the birds may be wing-clipped. To inhibit constant tra\elin2 up and down along the sides of the pen, called "walking of the wire"' it is wise to co\er the outside of the pen with burlap or other material for a distance of a foot above the floor. The feeding equipment may be placed in one corner under a shelter. If a few evergreen boughs are placed at one end the birds will s])end the greater ])roportion of their time in the co\er thus provided. T ihal later di'.-rribcd [or luinil-rai:-i'(l breeders pcinicrl al llic same season of the \car. Wild birds are mor<' likely to mate and la\ iliiririg llic s<'(()n(l ywv in rapli\il\. llowexer a certain ])rop(irlion will alwavs remain lam,' but nnpiiMlucti\e. Till. \ VTIKAL COVF.H METHOD or 1! \1SI\G CROLSE In adapting an\ species to ca|)ti\ ity. the first thought is to place it under conditions as nearly similar to its nati\e habitat as possible. It is small wonder that Clark ' in lo79, ])Ut four wild partridges, collected in llic fall, in a circular enclosure 10 feet in diameter with walls 5 feet high covered by a conical tcni. In il he set out close clumps of young pines and firs and covered the ground with dry forest lea\es. One of the birds subsequently nested and laid 18 eggs. Others, too. had the same idea. It will be remembered* that, to jirovide his breeders with natural wintering habitat. Hodge'"' some 30 years later completely co\ered a large spruce tree with poultrv netting in the front yard of his home. Two of the birds placed therein were snagged by cats through the inch-mesh wire of the enclosure and died of their wounds. The rest were poisoned. The American Game Piotective Association, in 1912. established a grouse farm on (]a])e Cod. and Torrev"' luiilt there a wintering and breeding pen of fish net- ting resembling a circus tent. This was placed so as to enclose a heavy growth of young pines. But disease and predation forced the abandonment of the idea after a year's trial. On the other hand, in Manitoba. Rendick'" for years sold grouse said to have been reared from eggs laid In wild birds held captive in a large enclosure. Between 1919 and 1929 Dr. Allen" repeatedly tried to raise young birds in pens containing natural cover. Breeders, too. were thus confined, but few escaped the ravages of "enteritis", jiredators and unfavorable weather conditions. Still not satisfied, the Investigation, in 1931, Iniilt a series of covered pens enclosing several * See Chapter I. p. 23. 454 ARTIFICIAL PROPAGATION OF GROUSE acres of excellent grouse cover. This was composed of pine, hemlock, laurel and hard- woods. Every effort was made to keep the pens disease-free. No poultry had ever pre- viously occupied the site and none was permitted close by. Yet, of the birds liberated therein, disease and predators took a heavy toll. Nor was egg production, fertility or the general condition of these birds as good as has been obtained with grouse held on wire. '•>' Gardiner Bump THE INTERIOR OF A PEN USED 10 H VISE CROL SE U\ THE SEMI-NATURAL METHOD AT THE CATSKILL STATION The first year 15 wild grouse were secured earlv in A])ril through the courtesy of Mr. R. W. Tufts of Nova Scotia. Two died in transit and ten were paired on the ground in 9 foot x 12 foot segments of the pens described above. The remaining three, together with two locally trapped birds, were liberated in the rest of the enclosure, totaling an acre and a half. Only one female nested, laying bul four eggs, all of which were infertile. A single bird, hand- raised the year before and confined in another ])en with a wild-trapped male, laid 25 eggs. Only three of these proved to be fertile. The second year, ten wild-trap|)cd grouse, placed in these pens in December and January, laid 41 eggs, 65 per cent of which were fertile. A similar number of hand-reared females pro- vided real encouragement by producing 165 eggs. 68 per cent of which were fertile. An indication that the ground possessed no uimsuallv stinudaling powers which could not likewise be brought to bear on birds held on wire came from Ithaca where Dr. Allen was experimenting with thus raising grouse. From one group of his breeders held on wire, 96 per cent of the eggs produced, proved to be fertile. Likewise, at the Research Center females have since laid up to ."^6 eggs each in a single year although their feet had never touched the ground. 'M With the above picture in mind, one may wonder why, on succeeding pages, the natural cover method of raising grouse is described at all. Though the difficulties inherent in it have engendered in the author a firm belief that no large number of grouse can be raised by this method over a period of years, a description is included for the sake of completeness and for those who are willing to chance the risks in this method. Furthermore, there are game breeders who still recommend it. THE NATURAL COVER METHOD OF RAISING GROUSE 455 Penning Birds and Providing Shelter A concensus of opinion dictates the choice of a well drained, fairly open site for all pens. The size of the wintering enclosure must of course vary with the number of birds to be held therein. From the standpoint of preventing disease there is no safe minimum, for a few birds liberated on an acre of ground are still likely to become infected. At the Catskill Experimental Station about 40 adults were wintered in a pen 54 feet x 75 feet x 8 feet without encountering serious losses. Subsequent experience, however, indicated clearly that one should expect an increase in mortality caused by blackhead after holding the grouse on the ground. Such a wintering pen should contain considerable natural shelter. Jeffries'™ claims that the proportion of shaded to sunlit ground is important and recommends that 60 to 75 per cent of the area is shaded. Sunlight may, however, be a strong aid in controlling disease. For this reason, at the Catskill Station the wintering and breeding area was left but one-quarter shaded. In this portion the birds tended to gather, and a wire floor was constructed over that part of the ground to reduce the danger of infection. The pen should be constructed of sturdy material and covered with 1-inrh poultry net- ting strongly supported to withstand winter snowfall and to discourage predators. At the Catskill Station, where 2-inch mesh was used, in one night a great horned owl beheaded five grouse. Frightened by the air raid, they had persisted in attempting to fly through the top of the pen until beheaded. To prevent the breeders from "walking the wire", boards or burlap about 12 inches high may be placed on edge along the side walls next to the ground. In February or early March the breeders may be placed, preferably in pairs, in small enclosures. The "all-purpose" pen (figure 37) is ideal for this purpose. By covering the windward end of the pen, shelter against inclement weather for adults and chicks alike, can be secured. In the spring, breeding females need small hideaways where they may seek pro- tection from the male. In fact, if the female is to incubate her own eggs, the males should be removed from the pens when the clutch is completed. Many suggestions detailed later in the chapter will be found applicable to the care on the 456 ARTIFICIAL PROPAGATION OF GROUSE ground of old and young alike. For instance the incubation of the eggs collected from the penned birds is fully described in a separate section devoted to this subject (p. 473 I. If one is also determined to try raising the young grouse in contact with the soil, the pens should be small enough to be moved easily to a fresh site at frequent intervals. Of course they should be kept scrupulously clean. One of the "all-purpose" pens placed on the ground contains plenty of room for a single brood. Particular care must be taken to chink all crannies where the pen meets the ground for the birds are adept at squeezing through small openings. In fact, when first hatched, thev have no difficulty in slipping through inch-niesh poultry netting, so that %-iiich mesh must be used. Feeding The literature is rich in feeding suggestions, many of which are impractical because the material recommended is not easily available. Ant eggs, bee larvae, maggots and mealworms are interesting examples. At the Catskill Station several tons of beef lungs were consigned to the "niaggoteria" every year. The lungs were divided, placed in milk pans and exposed to the sun. Later the pans were placed in a beetle-proof cage until the larvae, now fully grown, migrated over the edge into other pans containing bran. Here thev quicklv clpuiied them- selves and were ready for use. There was always the danger of botulism with this method. Accordingly, in 1931. a grad- uate student at Cornell was commissioned to develop a method of producing large numbers of fly larvae that feed on plants. A vear later, however, satisfactorv substitutes for fly larvae were discovered and bf)th the ])raitice and the experiment were discontinued. Although there is still sonic controversy among grouse breeders concerning whether best results may be obtained by feeding natural food, the difTiculty of obtaining it in large quan- tities has dictated the use of the feeding formulas and techniques later described for birds held on wire. Protection from Predators Birds held on the groiiiul arc notably exposed to attacks li\ jiredators. In late August 933. at the Research Center. 40 wing-dipped breeders were ])laced in an open jien several acres in extent. The enclosure was located near the middle of a meadow and was at least 750 feet from any woodland. Yet the first night a great horned owl, later trapped on its kill, decapitated six indixiduals. A somewhat similar case has previously been mentioned. On another occasion young grouse, wing-clipped and placed in a pen similarly situated, were promptly killed. The evidence pointed to a Cooper's hawk as the responsible predator. Birds on the ground are also susceptible to attack by rats. cats, weasels and dogs. On one occasion two dogs forced the entrance to a ])eii at the Research Center, killing 14 nearly grown birds. Raccoons arc also occasionally destructive. A grouse hen with her brood, placed in a wooded cnrlii~iii('. was destroyed three nights later by a "coon whose subsequent depredations were cfTcdixcK cliccked bv placing an electric fetice along the to|) wire of the pen. The best |)ri)tc(tii)M i> to Imild liic |)cii> tiglitK. idvcririj: llicrii u illi mikiII nioii wire and to place tunnel traps at frequent intervals along the outside boundaries for weasels and rats. These may be screened with two-inch poultr\ netting to prevent the entrance of escaped birds. THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 457 Danger from Disease Disease difficulties have caused the termination of many a grouse experiment, though, contrary to general belief, grouse are probably no more susceptible to this trouble than are bobwhite quail. As previously mentioned, to place grouse on the ground in confinement is to invite disease. To keep this to a minimum, feed and water should be placed on wire- covered frames and the surplus frequently removed. Any accumulations of droppings, par- ticularly under shelters, should be regularly cleaned out. A wire bottom built to fit each breeding or rearing pen is a wise precaution against a possible outbreak of disease. Pens and birds should be placed thereon at the first sign of trouble. Though the records contain a few instances where broods of grouse were reared more or less successfully with bantam hens, as Dr. Allen aptly put it. "One might almost believe the successful grouse breeder should not eat poultry even on Sunday", so susceptible are the birds to several of the more widely spread chicken ailments. Nor are fresh ground and seclusion guarantees against disease. As already mentioned, birds both young and old died from blackhead or from ulcerative enteritis fquail disease) in the large covered pens at the Catskill Station, even though great care was taken to avoid contamination and no poultry had previously occupied the site. Another year, at the Research Center, 75 breeders, all raised on wire, were divided among 15 pens placed on fresh ground. Extreme care was taken not to introduce disease, yet 16 of these birds died from lilackhead before the breeding experiments were concluded. These experiences, verified In those of other breeders, point inescapably to the conclusion that no large number of grouse can be raised or carried in ca|)tivily in contact with the ground without sooner or later suffering a high loss of birds from disease. Vitality The question is often raised as to the vitality of generations of grouse raised on wire in comparison with those reared and held on the ground. The situation is, in fact, purely aca- demic, for few if any breeders have been able to avoid the pitfalls long enough to bring captive grouse beyond the second generation on the ground. Neither do wild birds offer a valid comparison. In contrast, grouse have been raised and maintained on wire at the Research Center to the eleventh generation without experiencing any of the much feared degeneration of stock. THE INCUBATOR-BROODER METHOD OF RAISING GROUSE Six thousand years ago an inventive Egyption modeled in mud and baked in the sun- light a queer structure for the purpose of hatching out the eggs of the half-wild fowl that fre- quented his dooryard. Perhaps his friends laughed at the contrivance but it worked and eventually better ones were built. Thus out of an interest in the harnessing of nature's forces to man's ends was born the artificial method of propagating the birds of field and forest. At first game breeders were much prejudiced against applying artificial methods to the raising of game birds. But the comparative freedom from the destructive forces of weather, predation and disease thus afforded has forced an increasing number to adopt them during the past 25 years. Early results left much to be desired but refinements in methods and techniques followed rapidly after the discovery that the new way was markedly less expensive than the old. 458 ARTIFICIAL PROPAGATION OF GROUSE With a species as susceptible to disease as is the rufled grouse or the hobwhite quail, the choice seems not to lie between rearing methods. Rather it is dependent on how far one can go in applying artificial techniques. Generally speaking, quail and grouse are alike in that each can adapt itself rather successfully to meet most of the limitations of being kept on wire. r ! -i--^' .J- '",JS*- -"?'°^ - - - - >.v„^ >■■ I •:-■■■ W'"' EARLY EGYPTIAN INCUBATOR A glance at table 68 will indicate that the differences which make game farm production of quail much more successful than grouse propagation exist chiefly in the differential in egg production and in the rearing success with the young birds. TAHI,K 6!!. PENNING AND PUODtlCTIVITN OK BOBWllITK QUAIL COMI'MIED W 11 II RUFFHO (iliol SK IN CAPTIVITY Penning or productivity Quail Grouse Mrlliod *»f penning breeders. . . Pairs Pairs Size of pLMis needed 4' X 8' J :>' 6' X 8' X 3' - 8' X 16' X .T Normal Kgg production 50- 80 15-25 85 - 90% 90% 75 - 80% 12% 65 - 80% 70 - 80% 30 - 50% 15% Normul overwintering losses. . . To those who would raise grouse i>y the inculiator-broodcr method, a detailed considera- tion of the methods employed and the attendant results should be of interest. In presenting this picture, each natural unit of operation, such as breeding, brooding or rearing, together with its attendant problems and practices, such as feeding and preventing disease, has been handled as a suljject complete in itself. Only by so doing can one secure a well integrated idea of the way in which each period or problem can be handled. THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 459 Caring for the Breeding Stock One key to the successful production of any game species is the quality of the breeding stock*. With grouse this is all-important, for the gradual increase in the number of fertile eggs laid per female through selective breeding is a prerequisite to large-scale production. Once secured, there are three problem periods to be considered in handling grouse breeders. Least attention is usually given to the over-wintering of the breeders. This is a mistake for it is then that the number of eggs to be produced the following season is largely determined. Most attention is paid to the spring period when the breeders are actually mating and laying their eggs. Likewise, there is a tendency to forget the adults again while they are moulting and recuperating during the succeeding summer and fall. Over-Wintering Grouse Breeders Pens and Penning. By the first of October the birds of the year become restless in their rearing pens. Adult breeders have completed their moult and are ready to be moved into winter quarters. Although it is advisable to separate the males from the females, larger numbers of birds can be held together throughout the winter than at any other time. Adult grouse are not particularly disturbed by normal rains, snow or sleet. On sub-zero Gardiner Bump \\li\TKKli\(. \AKDS AT TlIK KKSKAKCll CKMLK C.O.MAlMiNG ALL-l'LUrusL TEiNS I'LACED END TO END days their movements are restricted somewhat but they do not appear to be uncomfort- able. The one item of weather which they dislike particularly is wind. Windy days are nerv- ous days for grouse both penned and in the wild. At the Research Center, two all-purpose pens placed end to end, making a single enclosure 8 feet X 32 feet x 3 feet with a shelter at either end, easily house up to 20 breeders. Eight birds can be fairly successfully maintained over winter in half this space, though the work of caring for a smaller group would, of course, be doubled. To provide perches and places * For methods of obtaining breeding stock, see p. 451. 460 ARTIFICIAL PROPAGATION OF GROUSE o o a u u ? 71 a z O OS < ca z" o z S b) H Z X H H M « THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 461 r THE WINTERING PEN l.s DIVIDED iiNTU SECTIONS B^ LOW BOARDS TO DISCOUKACE THE BIRDS FROM CHASING ONE ANOTHER THE SERVICE ROOM IS CONSTRUCTED WITH SLIDING WINDOWS AND DOORS THAT OPEN INWARDS TO FACILITATE CATCHING, FEEDING AND WATERING THE BIRDS OUTSIDE 462 ARTIFICIAL PROPAGATION OF GROUSE of escape needed by all save the dominant bird in each pen, two or three small pines or hemlocks may be cut and stacked butt up in a convenient corner. Allen found that a series of boards each 10 inches high and placed on edge and crosswise at 4-foot intervals helped to maintain the established pen society by permitting the setting up of several territories there- in over which individual birds might exercise dominance. This materially increases the chances that each bird in the pen will find among the several communities thus established, one or another into which it can fit. To many this may seem like drawing a fine distinction, for the birds, of course, do not always keep to the particular section of the pen where they find their companions and sur- roundings most congenial. Nevertheless, when chased, their own part of tlie pen offers a haven into which a combative outsider is often loath to continue pursuit. The matter of dominance, as discussed in Chapter II, is of considerable importance, for each year adults of both sexes are lost as breeders through being harried by more aggressive birds as to acquire an inferiority complex. Some birds are killed outright in the process and many are severely scalped in attempting to escape. The size of the over-wintering loss depends largely on maintaining the social balance in the pens. Of course there will also be an occa- sional death from accident or from chronic diseases such as tuberculosis, blackhead or ulcera- tive enteritis. A wired-in wintering pen 25 feet x 110 feet built around a service room (figure 33) and designed to hold 300 full-winged grouse comfortably, was put in operation at the Research Center in 1937. Here each bird has a relatively large choice of companions and of com- munities. Heavy cotton netting is stretched along the sides to ease the impact if a flying bird should misjudge distances. Most grouse quickly learn to alight before reaching the netting. The daily flights up and down the pen seem to furnish a good conditioning routine. Foods and Feeding Techniques. The food, too, during the overwintering period is im- portant, for the young birds have just completed the strain of growing up and the adults the stress of moulting. Each must build up resistance and reproductive capacity for the breed- ing season to come. Nor is it necessary to resort to special foods augmented by a variety of the items wild grouse normally consume at this time of year. The composition of the feeding formula currently (1942) in use at the Research Center is indicated in figure 41. All of these may be purchased at reasonable prices from many commercial feed concerns. Mash, in pelleted form, and grain are fed in low, reel-type troughs placed so as to be protected from the weather. In the wintering pen at the Center all feeding and servicing, as well as the catching of the birds, is carried on from the central service room by means of feed shelves and sliding win- dows. Water is provided by automatic fountains of one-gallon capacity set on thermostati- cally controlled electric plates to prevent freezing. In the smaller pens where electricity is not available, earthenware dishes, set slightly above the level of the wire, will suffice. These must be filled daily and cleaned frequently. The lengthening days of early March bring with them increased activity and physiological prejiaration for the breeding season. The birds, whose consumption of corn in winter will vary with the temperature, now pay little attention to this grain. Since weight is a factor in egg production, it is wise to tempt their appetites further by setting before them a variety of cereals. A moist mixture containing three parts of laying mash to one part of cereals and the substitution of lettuce heads for dried alfalfa, assist in accomplishing tliis purpose. These THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 463 changes should be made at least a week before the birds are placed in the breeding pens, thus reducing somewhat the confusion incident to the change. Although the ration here suggested seems to meet the needs of the birds rather well, it is desirable to provide them with supplemental foods, such as apples, and alfalfa hay, in a rack. These serve to keep the birds occupied and accordingly better satisfied with their quarters. It is interesting to note that some difficulty may be experienced in inducing grouse to eat an unfamiliar food such as beechnuts or buds, even though it may be a staple diet of their wild cousins. Handling the Spring Shuffle. As spring approaches one may notice a gradual increasing unrest among the captive birds. The males strut with increasing frequency on the warmer, quieter days. In the wild this restiveness marks the beginning of the spring shuffle. Its counterpart in the wintering pen calls for close attention to the dominant birds, for they are most likely to scalp or kill their inferiority-coniplexed companions at this time. It may be necessary to segregate many of the more active males until the birds are placed in the breed- ing pens. Such separation is undesirable if it can be avoided for these males must reestablish their social position in the flock when they are returned*. In the latitude of Albany it is wise to pair off the birds about the first of March, utilizing the suggestions for the selection of breeding stock as described later. Weight and Physical W ell-J>eing. In grouse, as in most other birds, weight is a fair indi- cation of physical well-being. Chronically diseased individuals often may be discovered if the birds are periodically weighed and the results checked with table 69. TABLE 69. AVERAGE WEItillTS l.\ GHAMS IJF HAND-REARED GROUSE AT THE RESEARCH CENTER DURING CERTAIN PERIODS Age and Sex I'ericKl Over oue year old Birds of the year Male Female Male Female Nov. 15th March 15th June 15lh . ... 625.6 5>)0.2 60U.0 565..1 578.9 502.3 S'):i.8 586.1 551.6 543.5 516.7 511 5 The Spring Period Selection of the Breeding Stock. The success of artificial grouse propagation lies to no small extent in the quality and condition of the breeding stock, for this is the foundation upon which the year's work is built. The qualities inherent in these birds, coupled with proper care and attention, are reflected in egg production and hatchability and in the subsequent brooding and rearing success. Results clearly indicate the need of a selective breeding program in order to develop a strain of grouse possessing as many desirable characteristics as possible. Space does not per- mit more than a brief resume of the breeding plan followed at the Research Center. While most persons interested in improving their own strain of grouse may want to develop their own plan, this synopsis might be useful as a guide. The program is designed to encourage » See Chapter II. p. 63. 464 ARTIFICIAL PROPAGATION OF GROUSE the establishing of groups or famihes of grouse capable of high egg production, hatchability and livability. The method of establishing; these families is an important consideration. Beginning in 1935 all females were classified into four groups depending upon whether they laid in excess of 25 eggs in a single season, from 20 to 24, from 15 to 19, or below 15. In each of these groups the females were further rated according to the percentage of their eggs which were fertile, the percentage which hatched and the number of chicks which subsequently survived. Each male was rated on the basis of his parents' and his own performance records and on his ability to get along with his penmates. All these items were carefuUv considered in mating the birds each year. As a result, family groups, containing the characteristics desired, were built up. Birds from these groups are the foundation of the breeding stock. Progeny testing or analyzing the performance of the offspring of each pair of birds, has disclosed errors in previous matings and provided clues for correction. While the selective breeding plan has not be?n in effect long enough to disclose any far reaching results, it is evident that such a program may go far towards answering many prob- lems. Progress has been necessarily slow since it is difficult to bring through an adequate number of the progeny owing to the high mortality during the brooding period. It is essential in such a program to maintain a complete record (figures 34 to 36) of pedigree and performance covering the history of every bird. This begins with the breeders, both male and female and includes the past performance covering egg production, fertility, hatchability and livability, not only of the individual bird but also of its brothers, sisters and ancestors. There is another factor which should be considered. In addition to choosing birds from families with outstanding performance records, it is also desirable to select for type and temperament so as to retain the wild characteristics. Size, weight, structure and general body conformations are other important considerations. It would seem a good practice to main- tain the average for wild birds. For example, it would ])rolialil\ be unwise to attempt to improve upon nature to the extent of producing a strain of grouse all wcigiiing two ])ounds or over. The birds retained for breeders should be alert and thriftv in appearance, ullli llu- funda- mental characteristics of a wild bird. Those with any physical abnormalities such as long or short necks and legs as well as those heavier or lighter in weight than normal might bettor be discarded. After the breeders ha\<- been selected and a decision nuulc in regard to lln' matings. they are placed in tlic breeding pens. lirecdinii Pens and F(/uipnienl. To date there seems to have been no grouse breeding pen developed which has been accepted as standard. There are, however, certain prerequisites which might well be considered. When dealing with large numbers of birds, it is important to construct and arrange the pens so thev may be easilv servi<-ed. Likewise, those which are largely self-cleaning and convenient to keep free of debris and to disinfect arc. of course. preferable. Adequate floor space for the normal activity of the birds is essential to general hi-alth and well-being. While the grouse appreciate some shelter from the elements, it is desirable to admit sunlight to a portion of the pen. To prevent aimoyance by the male, par- THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 465 fa 11 u 1 £ £ £ & £ c > 4- L 1 > C L. 1 > b c 1. > * » > 1. ■E C * 1. ■. > b 211 in Q a a O O w o ^1 h. a & & & ■ t I? en Q Z a o C S> 5> >N' OG^^ r IS 1 Daughter uumbor ;S 1. •al 9 a i ll 5" 1 B-2 ll 15 1 a OS o (J u » U b: u S u 0. o z w « o fa oe u H z u u s o u K U X H ■< O Id S o fa CO u a: 3 o o LIBRARY l» ^ MASS. /^J 466 Number of Male. Date Hatehed ARTIFICIAL PROPAGATION OF GROUSE INDIVIDUAL MALE RECORD I)nri-ct«: Date of death. . Cause of death . Year Mat£d to hen number Number of eggs set Per cent of eggs infertile Per cent of fertile eggs hatched Number of chicks hatched Number of chicks rearwl to maturity Remarks Male Female FIGURE 35. FORM USED FOR KEEPING RECORDS OF INDIVIDUAL MALE BIRDS INDIVIDUAL HEN RECORD Number of Ilnn. Date Hutoheil. . . DcfRCtJs: Date of death. . Cause of death. Year Pen Production Date of 1st egg Date of last egg Remarks Year Mated to male number Number "'■air Per cent of eg^ infertile Per coat of fertile eggs hatched Number of chicks hatchoniarks Male Female FIGURE 36. FORM USED FOR KEEPING RECORDS OF INDIVIDUAL FEMALE BIRDS THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 467 ticularly during the breeding season, protective cover should be provided for the female. It it sometimes difKcult to incorporate all of these factors into a single pen without running into considerable expense. The number of birds that are to be accommodated will, in large measure, determine the basic design and arrangement. Only by trial and error will the type best suited to a particular purpose be developed. Many sizes of pens, mostly of the coop and run type, have been tested experimentally at the Research Center. The 8 feet x 16 feet x 3 feet all-purpose pen (figure 37) proved quite satisfac- tory when polygamous matings were being tested but was found to be larger than necessary when the birds were mated in pairs. A pen measuring 8 feet x 12 feet x 3 feet was accord- ingly designed to accommodate such a grouping. This met most of the requirements and was used as the basis for the development of less expensive, permanent, multiple-unit breed- ing pens. It was later found that the floor area for a pair of birds could be reduced further if certain modifications were incorporated. With this in mind four such pens were constructed and each subdivided to accommodate 12 pairs of breeding birds. The over-all dimensions are 8 feet X 72 feet. The pen is built on posts with the floor 2V2 feet above the ground. The individual unit housing each pair of birds is 6 feet in width by 8 feet in length and 3 feet in height. One side of the pen is enclosed and roofed, making a coop shelter measuring 2 feet X 6 feet in each individual breeding compartment. The rest of the pen is enclosed with %-inch mesh poultry netting, the sides and top being 20 gauge. The floor under both coop and run is two gauges heavier. Each run is separated from the next by a removable parti- tion which slides upon a 12-inch baseboard. Kach coo]) j)ortion is separated by a hinged door. All |)artiti()ns may be removed at the end of the breeding season, allowing the birds more freedom. The males may be segregated in one half and the females in the other. Feeding equipment, after many trials, was selected from among the several types manu- factured for game birds. For a pair of grouse, two low 12-inch metal feeders are satisfac- tory, one for grain, the other for mash. Water is furnished in the multiple-unit ])('ns through automatic fountains while, in the other pens, clay water dishes 4 inches in diameter and 1 inch deep are used. The same type of dish may be used for feeding wet mash. Feeding and watering equipment and a removable, open-type nest box are sheltered in the coop and serviced through doors conveniently located. The nest box, 12 inches square and 6 inches high, is lined with grass. This must be replenished at intervals since the birds will consume most of it. It is interesting to note that the built-in nest boxes in the all-purpose pen did not seem to meet the full requirements of the birds. These boxes were totally enclosed, the birds enter- ing through an opening in the side. The females seemed more contented when the open-type nest box was used. This appears logical since the wild grouse, which nests on the ground, normally selects a site which is open from above. Penning and Care of Breeders. After the pens at the Center have been thoroughly cleaned and disinfected and are ready for occupancy, the breeders are transferred to them. One pair is placed in each individual unit. Fresh evergreens are placed in the corners of the run to give the birds seclusion. With the coming of spring, there are certain social readjustments to be made by the birds. 468 ARTIFICIAL PROPAGATION OF GROUSE The males will be seen more frequently in their courtshij) display. At the Center the birds are moved into the breeding pens early, usually by the first of March. This encourages harmony among the penned pairs and results in an increase in the number of fertile eggs |)rodualirtant physical factors of incubation — temperature, humidity and THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 477 air movement. During the study the basic requirements for artificially incubating pheasant and quail eggs were determined and those for grouse eggs were outlined as a result of experiments with some 350 eggs. "still air" incubator with turning trays to hold incubating eggs and pedigree trays into which the eggs are shifted before hatching Beginning in 1933, a series of tests covering incubators and incubator conditions were also carried out at the Research Center where incubator techniques were set up, carefully checked and modified as appeared desirable. Laboratory recommendations, resulting from the Cornell experiments, were also tested on large numbers of grouse eggs under practical condi- tions. Thus was secured the background for the recommendations detailed later. The studies carried out to date indicate that grouse eggs may be incubated and hatched more successfully in properly constructed still-air incubators than in agitated-air machines. No unusual situations or difficulties were encountered in securing successful hatches of eggs in the former type. In general the operation of the machine is not materially different from that employed with pheasants or quail. The conditions recommended in table 70 will serve as a guide for the operation of the incubator. 478 ARTIFICIAL PROPAGATION OF GROUSE TABLE 70. SUGGESTED CONDITIONS FOR THE INCUBATION OF GROUSE EGGS Types of incubators Incubatiufc conditions Still-air (hot-air or hot-water) Agitnted-air (with separate hatcher) 1st week 2nd-3rd week From 20th day Isl week 2nd week on From 20th day 103° F. 60-65% >4 open 3 to 4 times 103° F. 60-65% H-Ji open 3 to 4 times 103° F. 60-65% much rediK-t'd none 99>A° F. 60-65% 87-88° F. small 3 to 4 times 99)^° F. 60-55% 87-88° F. niixlcralc 3 to 4 times 99 H° F- ir. ™- J -.../relative Humidity^ ^^^j,„,h Vontilalion 60-657o »7-88» F large none In connection with the above table there are certain qualifications to bear in mind. Because of different rates of air movement in various makes of still-air type incubators the average reading of temperature may vaj;y from 102°F. to 104°F. The references given above are for a room temperature of 60°F. If the room temperature is low, for example 40°F., the incu- bator temperature should read about V2°F- higher; and, if the room temperature is high, for example 80°F., the incubator temperature should read about •!/2°F. lower throughout. This is to compensate for the current of warmer air entering through the ventilators. The bulb of the thermometer normally is placed about two inches from the egg tray level. The temperature should be higher when the bulb is more than two inches above the tray level and it should be lower when the bulb is elevated less than two inches. Because it is important to secure the exact temperature, it often is desirable to take a read- ing on clinical thermometers, the one placed on the floor of the egg tray, the other level with the top of the egg. The optimum temperature for grouse egg incubation is 99V^°F. at a point one-half to two-thirds above the bottom of the egg. It is a good practice thus to check the incubating temperature at the beginning of each successive hatch for, as the season advances, the temperature of the incubator room likewise usually increases. Unless one is skilled in still-air incubator operation, it perhaps is safest to open the venti- lators a quarter during the first week, one-half the second week and three-quarters from the 15th to the 20th day. After that the ventilation should be reduced markedly. Such sugges- tions of necessity are general and subject to modifications as experience directs. Whenever an increase in humidity is desirable, particularly at hatching time, all moisture pans should be filled to allow as great an area as possible for evaporation. Otherwise, they should be kept partially covered or left dry according to the humidity in the room and the condition of the eggs as indicated by the last recorded per cent of weight loss or by the mois- ture gauge. It is well to remember, too, that excessively evaporated or old eggs require a higher humidity than do unevaporated or fresh eggs. Mcasuririf^ Inruhalion Proprcss. As proficiency is attained, a need is sensed for more def- inite methods of determining the progress of the hatch. Two techniques are helpful — peri- odic candling of the eggs in order to check on the progress of development and tlie checking of weight losses to determiiK' the rate of evaporation of moislurc from the eggs. When one becomes skillful in interpreting these indices, they will provide many a cue for modifications of the suggestions heretofore mentioned. For instance, too rapid development of the embryo itulicalps that the iiuubalor is being run at too high a temperature. This must be lowered until the normal schedule of development is resumed, else dead germs and poor hatches are THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 479 likely to result. If the eggs have not lost sufficient moisture at a given point in incubation, it is necessary to reduce the humidity and increase the ventilation to permit more rapid dry- ing through the shells. In the still-air incubators temperature also is affected by ventilation changes and, thereafter, must be readjusted. Although the shell of the grouse egg is almost as thick as a pheasant's, averaging 0.21 mm. it has less pigment and is less opaque. It is wise to candle all newly set eggs on the third day. As pointed out before, those that are infertile may then be identified although not removed and arrangements made immediately to place a different male in the breeding pen in which the eggs were laid. Fresh egg Inlertile egg, ftlh day Fertile egg, .ith day Dead genu after Ith day FIGURE 38. APPEARANCE, UNDER THE CAMJLIiNG LAMP. OF FERTILE AND INFERTILE EGGS DURING INCUBATION A little experience, aided by a careful study of figure 38 will assist the beginner to detect the clear eggs which are infertile. As incubation progresses a live embryo may be identified under the candling lamp as a dark floating spot from which blood vessels radiate. Under normal development, by the 12th day the blood vessels have advanced until they meet at the small end of the egg. Just prior to hatching, the egg will be opaque except for the area occu- pied by the air cell. Once one has become skilled in recognizing these changes, it is helpful to candle the eggs on the 12th day and again on the 21st day, just before shifting them to the hatching compartment. The practice of checking weight losses to determine moisture content is also distinctly helpful, for unless a chick hatches clean it is apt to die or be weaker than the rest. Careful experiments at the Center have indicated that grouse eggs hatch best when about 13 per cent of their weight is lost in the first 21 days of incubation. The following is a simple formula for determining the percentage of weight loss at hatch- ing time provided the existing rate of moisture loss is maintained. WL WE X " X 100 = Percentage loss in weight projected to hatching time WL = Weight loss at the time of weighing WE = Weight of eggs at time of setting I = Incubation period (231/^ days for grouse) D = Number of days eggs have been incubated at the time of weighing 480 ARTIFICIAL PROPAGATION OF GROUSF The figures needed to apply this formula may he easily secured. By weighing the empty Iray and the tray with eggs on it at the time of setting and hy suhtracting the former from the latter, the weight of the eggs at setting time is determined. The weight loss at any subse- quent point in incubation may be found by similarly finding the weight of the eggs at the desired time and subtracting it from the weight at the lime of setting. Many factors, among them temperature, ventilation and outside atmospheric changes, in- fluence the regulation of humidity. Obviously, even though a humidity curve such as tlie one illustrated may represent the optimum, a successful hatch of grouse eggs will not occur unless the other factors are also properly balanced. The correct weight loss, hypothetically. would be represented by a straight line. However, owing to the influence of these other factors, such a line rarely exists. In order to visualize the rate of moisture loss, figure 39 may be helpful since this weight curve represents the moisture loss in a setting of grouse eggs of which 87 per cent hatched. I/) o I o 12 13 14 DAYS INCUBATED FIGURE 39. PROGRESSIVE MOISTURE LOSS DURING INCIBATION OF GROUSE EGGS Since the early removal of infertile eggs or those with dead germs would complicate the com- putation of weight loss, they arc left in the trays until such determinations are complete. Selling and Care of Eggs. While it is best to set the eggs within a few hours after they are collected, they may be stored without undue risk in trays, preferably at temperatures between 40° F. and .SO" F.. since incubation commences at about 60° F. To keep the germ spot or blastoderm from sticking to the shell, the eggs should be turned at least once a day. A tray THE INCV BATOR-BROODER METHOD OF RAISING GROUSE 481 with an automatic turning device, as commonly used for pheasant eggs, is satisfactory for this purpose. Because of probable deterioration, it is not advisable to hold the eggs longer than ten days before setting. Those from wild nests, if they are already partly incubated, obviously should be placed in the machine at once. Dirty eggs should be scraped with a penknife or wiped with a damp cloth before storing or setting, for washing destroys shell cuticle. Misshapen eggs, as well as those with badly cracked shells, should be culled out. Those with minor cracks will sometimes hatch if the in- jured area is covered with shellac. Eggs in the incubator tray should never be placed with the small or pointed end up lest one encourage embryonic deformities. The normal incubation period for grouse eggs is 23V2 days. Occasionally, because of undue heat, a few may hatch in 23 days. In the wild, if the weather is cold, the hatching date may be deferred until the 25lh or even the 26th day. The wild grouse mother may leave her nest on warm days for several hours at a time. But at the Research Center care is taken to avoid chilling of the eggs during incubation ex- cept when required in carrying out candling or weighing procedures. Hatching and Transfer of Chicks. When the developing chick breaks into the air cell, usually about the 22nd day of incubation, it is time to shift the eggs into the hatching com- partment of the incubator. A separate section is utilized for this purpose. The eggs from each female are placed in individual compartments or wire baskets called pedigree cases so that the identity of each chick will not be lost. There are several successful ways of handling the eggs at hatching time. At the Center, as soon as the first egg pips, the ventilation is reduced to conserve humidity and the moisture pans are often filled with lukewarm water. If the chicks pip the shells but have difficulty in getting out because of a toughened egg membrane, they may be sprinkled with tepid water. Doug Fincit CHICKS MAY BE IDENTIFIED BY ATTACHING A BAND TO LEG OR WING BEFORE TRANSFERRING THEM FROM THE PEDIGREE HATCHING TRAY TO THE BROODER 482 ARTIFICIAL PROPAGATION OF GROUSE Do not open the incubator more than is necessary during this period. If egg development and incubation conditions are right, the hatch should be complete within 18 hours of the time the first egg is pipped. The new arrivals do not require food or water until they have been transferred to the brooder for, like the camel's hump, nature has provided the chick with a large unabsorbed yolk in which much of the food for the first two or three days is stored. When the chicks are completely dry they may be weighed, wing-banded and placed in a small clean box, the bottom of which has been covered by a soft material, such as cloth. If the weather is cold, the box may be warmed and wrapped in a blanket to make certain that the chicks do not become chilled in transit to the brooder. Brooding of the Chicks In any extensive wildlife research project, one or two ba£Bing problems are to be found which common sense tells us can be solved but which may elude the best efforts of the re- search worker for many years. So it is with the low rate of survival during the brooding period. The largest loss experienced in raising grouse still occurs before the chicks are four weeks old. The problem is puzzling, for chicks held under apparently identical conditions and fed the same ration may thrive or die without apparent reason. To all appearances the same con- ditions that encourage the survival of 60 per cent of the chicks started one year will result in but half that number pulling through the next. Many of the dead birds seem to be in good physical condition and without evidence of disease. Following the lead of earlier experimentors, the Investigation carried out exhaustive tests of brooders and brooder houses, of hover operation and of all the various other items so important in the care of chicks. Feeds and feeding techniques were repeatedly checked and changed with the aid and advice of collaborators skilled in the solution of nutritional diffi- culties. For two years Dr. L. C. Norris of the New York State College of Agriculture carried out reconnaissance studies of nutritional requirements of grouse chicks for the In- vestigation. Great progress has been made in reducing certain recognizable sources of loss, such as failure of the chicks to start feeding promptly, lung inllammation brought about by chilling and nutritional disorders due to improper feed. Other difliculties, notably feather picking and cannibalism and the danger from disease were also among those to which sat- isfactory answers were secured. But the underlying combination of causes of tliis early mor- tality remains the largest unsolved and disconcerting mystery in the field of artificial grouse propagation today. A dozen new methods have given high promise the first year in which they w^ere tried, only to prove stubbornly unreliable in succcrdinp (rials. Rcnicnibcrinp that in the wild the normal mother grouse loses up to 30 per cent of her brood during the first two weeks after hatching, it has even been suggested that grouse may possess a lethal gene similar to that recognized in some strains of poultry. It is axiomatic in brooding grouse chicks that the chances of success tend to he inversely proportional to the number of birds cared for. Ofltimes a single brood is raised with the loss of only one or two individuals. The answer, as far as is known today, lies in unremitting care and attention to the de- tails incident to feeding and brooding the chicks. Visitors at the Center have sometimes de- THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 483 scribed this situation as "fussing with the birds." Yet it is true that, even at this early age, each chick exhibits an individuality all its own. Game breeders, used to raising pheasants and quail en masse, may shake their heads at the thought of paying separate attention to the individual requirements thus discovered. But to one intent on raising grouse, nevertheless, it will pay big dividends in terms of chick survival. The more one works with grouse the dearer one realizes the necessity of paying attention to such minute details if one is to divert the forces of untrammeled Nature to serve Man's ends. Later it may not be necessary, for SHORTLY M'TKR II \TCHIN(. THE CHICKS ARE PLACED I\ A BATTERY BROODER selective breeding should produce a strain of grouse that can be raised in captivity, with proper care and without prohibitive early losses. Because of the obvious importance of the problems already mentioned, it should be helpful to treat each in some detail. Brooders and Brooder Houses As probably every game breeder knows, the type of brooder or brooder house influences (he method of caring for the chicks as well as the feeding techniques. The reverse also is true. Therefore the design of the housing facilities and equipment should be such that all of the activities incident to the handling of the birds will be expedited. 481 ARTIFICIAL PROPAGATIOX OF GROUSE 2 O o 13 > z o O as 3 THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 485 A decade ago it still was generally believed that nothing short of a broody hen could be trusted with the care of newly hatched grouse chicks. However, when a foster mother was used, the many failures largely due to the disease indicated a need for a mechanical-type brooder that could be kept clean. All the early types tested proved tricky and often unreliable. In spite of this, encouraging results often were secured. Allen gives a graphic picture of this situation in his "Ten Years Experiments in the Rearing of Ruffed Grouse in Captivity"". t-J*'^ x^ ■ ■r-ililiHiiV / A FLAT-TOPPED, BOX TYPE HOVER WAS FOUND MOST SATISFACTORY FOR BROODING YOUNG GROUSE IN THE COLONY BROODER HOUSE During the early years it was quite natural that attempts should be made to adapt the Cole- man type brooder, so successful with quail. At the Catskill Station 20 of these with their kerosene heaters provided enough work to keep the operator busy day and night. Some idea of the difficulties encountered is indicated when seven of these units caught fire and three were destroyed in one season. Herculean efforts were required to raise 46 grouse from the 189 which were started. The struggle required to maintain reasonably constant temperatures even with the best equipment soon prompted the search for a more efficient type of brooder. Considerable prog- 486 ARTIFICIAL PROPAGATION OF GROUSE ress was made by installing a small insulated brooding compartment in one section of the all- purpose pens already developed by the Investigation. In 1933 they were equipped with elec- triially heated hovers of sufficient size to accommodate 25 to 30 newly hatched chicks. Wire floors above removable pans to catch droppings reduced the chore of cleaning to the minimum. Other types were tried out in an attempt to find units that would accommodate still larger numbers of chicks. A circular brooder house, similar to that commonly used for domestic fowl and equipped with a standard .56-inch circular electric hover providing several heat zones, was tested. Several trials, utilizing from 50 to 200 chicks, were run with this unit. The results in all cases were extremely discouraging. It appeared that the proportion of losses mounted as the number of birds brooded together was increased. This also seemed to hold true in other types of units that were used. Not only was it impossible to give the birds the close attention which they received in small units but fluctuations in temperatures appeared to be an important factor resulting in high mortality. With these experiences in mind, a colony house was designed to overcome as many of the difficulties as possible. This unit. 12 feet x 100 feet x 10 feet, accommodates 16 individual pens. These are elevated so that the floor of the pens are waist-high for convenience in handling and to raise the chicks above floor drafts. The basal area of each pen is 40 square feet. An alleyway at the rear, as well as one between each set of two pens, facilitates the servicing and care of the birds. To exclude disease-carrying flies, fine mesh screening encloses the sides and sliding doors of each pen. An outside wire-floored run. also screened, approxi- mately equal in length and width to the interior pen but only two feet high, provides fresh air, sunlight and additional space for exercise for the growing chicks. To facilitate frequent removal of the droppings, tough, waxed wrapping paper fed from rolls is stretched beneath the wire floors of the run and above the floor screen. The exterior of this colony house is shown in figure 40. All the widely recommended hovers, including feather brooders, contact hovers, radiant agi- tated air types and others, were tested*. None seemed to give the close control that young grouse chicks apparently require. Thus it was that a standard liattery-type brooder was util- ized to house the chicks for the first ten days to two weeks. After manv tests, the most satisfactory brooder developed utilizes a heated compartment, enclosed on three sides, with a heavy denim curtain separating it from the run. This run is enclosed with tight-fitting removable wire mesh doors which are hinged at the bottom. With this arrangement there is no opportunity for lively young chicks to squeeze through openings or to hop out while the unit is being serviced. For easy cleaning, the whole can be disassem- bled by removing the sides, floors and dropping pans. The battery brooders are placed in a concrete-floored room with large windows affording an abiiiidanie of light. Room temperatures are thermostatically cf>ntrolled to remain as nearly as possible at 70° F. during the early spring. This serves to reduce temperature fluctuations in the batteries themselves. Brooder Operation To maintain condition" ideal for the growth and development of tlie chicks, much nuisl be left to the judgment nf (he Mllriuhuit. Rrotxling failures usually arc due either to faulty equipment such as conlrd" ulii
  • zziijOoa;tt;i-l->>oij ->-ii.-L.33<<33-)-iT=S<rtaiit to keep disease preventive practices constantlv in mind. Battery brooders should be thoroughly cleaned, scrubbed and disinfected before a new lot of chicks is started. The same applies to each unit of the colony house which has been previously occui)ied. Attendants responsible for the care of the adult grouse or of other species should, if possible, keep away from the battery brooder room as well as from other building or units where chicks are being reared. At the Center, as an added precaution, a mat soaked in disinfectant is placed before each outside door so that shoe soles can be dis- infected. THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 491 All feed for the young grouse should be separated from other feed. Likewise, there should be no exchange of feeding equipment or accessories unless they are adequately sterilized. Doors and windows in the battery brooder room and in the colony house are best fitted with tight screens to prevent the entrance of flies. The outside run, used in connection with the colony house, also should be screened. Control of mice and rats is important. The latter occasionally kill chicks. The former are not predators but their raiding of feed hoppers might possibly aid in spreading disease. Then, too, the unmistakable signs of mouse travels are left for the young chicks to pick up. Rearing of the Young BraDS As soon as the young grouse can do without artificial heat during cool nights, usually be- tween the fifth and sixth week, they may be transferred to the more spacious rearing pens. The brooding unit, while providing sufficient space for chicks, is costly to build if one makes it large enough to furnish the space desired from this time forward. At the onset of summer, birds are more likely to pick feathers or become cannibalistic. These, in fact, are the most likely causes of loss during this otherwise quiet growing period. Contrary to common belief this trouble may result as much from over-crowding or from fail- ure to keep the birds occupied, as from inadequate feed. Since grouse seldom develop a strong desire to dominate their penmates until the early fall they can be re-grouped in any desired combination in the rearing pens. During this adolescent period it is normal for tlie birds in tlie close social entity of the brood to devote most of their time to the business of growing up. In fact, at no other period, except when they are in the wintering pens, do the birds require less attention. The summer is, however, not without its dangers unless the requirements characteristic of it are properly understood. The type of rearing unit, the feed, the disease precautions and the recognition of the danger signals heralding the onset of unrest and aggressiveness in the young birds must all be considered if the rearing period is to pass without untoward difiS- culties. Rearing Fields and Units The ideal field, in which to place rearing pens, is a level, well drained site protected from strong winds and partially shaded. A properly situated apple orchard provides an excellent combination, for the trees may be judiciously trimmed all summer and the leafy branches placed in the pen to furnish satisfactory inexpensive roughage. Woodland glades have not proven satisfactory locations because of the number of grouse predators frequenting such locations and the difficulties of finding an occasional escaped bird in the surrounding under- growth. The field need not be large, for the rearing u.iits can be left permanently in the same loca- tion. To deter roving dogs, cats, rats and skunks and to keep curious visitors at a safe dis- tance, the area may be surrounded by a 6-foot poultry netting fence. The lower third should be %-inch mesh galvanized wire with the bottom anchored firmly in the soil to lessen access by rats. A few tunnel traps, properly screened with 2-inch poultry netting to discourage en- trance by escaped young birds, placed along the outside of the fence will catch many predators while they are following the fence. 492 ARTIFICIAL PROPAGATION OF GROUSE In spite of the preponderance of evidence against it, so attractive was the idea of raising the birds on the ground in large covered units containing natural cover, that three such structures were tried out during two rearing seasons at the Catskill Experimental Station. Although the birds were fed and watered on wire screens and the pens were kept meticulously clean to inhibit the onset of disease, severe outbreaks of ulcerative enteritis, coccidiosis and blackhead were encountered. Rats and weasels eluded every safeguard to take toll of the AI.L-l'l KPOSK I'ENS, .SCATIKKKII 0\ KU THK HKAK1.\(. HKl.I). MARK lUlCAL LMTS I.N WHU.H TO HAISK THE YOUNG CKOUSE TRANSFERRED FROM THE COLONY BROODER HOUSE young birds and great horned owls beheaded a number that, in fright, persisted in attempt- ing to fly through the top of the enclosure. This type of housing accordingly was abandoned in favor of much smaller units. Pens and Penning The prime requisites of a good rearing pen are spaciousness and accessibility. It should of course have a wire floor raised from the ground a distance sufficient to allow free access un- derneath for cleaning. It is desirable to provide a permanent shelter against unusual wind and rain at one end. Needed also are several roosts to which the birds may retire at night or take refuge when chased. To prevent "walking the wire," a 10-inch baseboard may be placed along all sides. The rest of the sides, top and base should be covered with smooth, galvanized "yi-inch poultry netting. Two types and four sizes of rearing pens were experimentally tested at the Center. Birds were raised to maturity in the colony brooder house, in 8 feet x 16 feet x 3 feet all-purpose pens and in units 8 feet x 12 feet x 3 feet and 4 feet x 8 feet x 3 feet of similar design. THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 493 The colony brooder units proved not too satisfactory because of the large proportion of enclosed house in comparison with the small run. It was found that the requirements of the older birds were best met by providing a minimum of 12 square feet of floor space per bird, in contrast with but four square feet needed by the younger chicks. As the birds grew older the periodic cleaning required in the brooder houses also was time consuming. iiarilintr Bump BIRDS MAY BE KEPT IN THE ALL-PURPOSE PENS IN THE REARING FIELD UNTIL WELL INTO THE FALL Of the three field rearing pens tried, only the smallest proved too limited in space. Under normal conditions, eight birds may be safely reared in the 8 feet x 12 feet x 3 feet unit, and ten to twelve in the larger one. If the pens are joined end to end and the intervening wire panels removed, still greater freedom of movement is provided, while maintaining the mo- bility of each unit. To provide shelter and to keep the birds busy, it is wise to lay small evergreens, butt up, in one corner of each pen. Rearing pens may be placed in rows at 8-foot intervals. A greater space between each pen sometimes facilitates the control of certain diseases but increases the time necessary to service each unit. Feed and Feeding Techniques The shifting of the birds from the brooder to the rearing field entails more than merely a change of pens. The moult from juvenile to adult plumage has already begun. 494 ARTIFICIAL PROPAGATION OF GROUSE III llie wild the young birds are paying less attention to insects and more to the leaves, seeds and fruits which largely make up the diet of adult grouse at this period. Likewise in the pens one finds an increasing tendency to eat more grain, in proportion to mash. The actual change in feed at this time is, however, not so much in composition as in size. The more coarsely-pelleted growing mash is substituted for starting mash, the intermediate-sized grains for the finer chick grains and intermediate grit for the smaller size previously used, as indicated in figure 41. By now the birds are also quite able to secure roughage by stripping apple leaves from the limbs. Lettuce may accordingly be eliminated and whole branches of apple or other edi- ble leaves placed in the pen. Beginning with the seventh week the birds will make increasing use of apples cut in half and impaled on nails. Bunches of clover or alfalfa, tied together and hung up or placed in a feeding rack from which thev may be pulled as needed, are also relished, though they wilt and pack down easily. These, together with needles from the ever- green boughs placed in a corner of the pen as shelter, not only offer the desired bulk to the diet, but also serve to keep the birds occupied. All substitutions and changes should be carried out gradually over a period of a week to cushion the shock possibly attendant upon sudden shifts in diet. To one who is raising grouse for the first time there comes an almost overpowering desire to add wild delicacies such as berries and insects to the standard diet here recommended. Yet sowbugs, for example, are known to be an intermediate host for the stomach worm (Dispharnyx) and grasshoppers may carry the gizzard worm (Cheilospirura). After yield- ing to this temptation for three years the Investigation discontinued the practice without any apparent adverse effect upon either the physical development, feather growth or the rate of survival among the growing birds. Little change in feeding or watering equipment is necessary at this time. Separate trough- type feeders for mash, for grain and for grit may be used, or the last two items may be mixed together. Small containers which require filling several times a week are preferable to the large hopper-type feeders. In the sununer moist feed should not be allowed to remain long in the feeders lest mold develop. Nor is it difTicult to supply the birds with water, reasonably fresh and cool. Feeling that the mason jar type of water fountain might encourage the spread of disease, a simple arrange- ment causing drops of water to form slowly at the ends of short glass tubes protruding into each pen, was developed in 1934. But while the birds enjoyed picking at the drops thus formed, the device required considerable servicing and was not easy to handle. Nor did it apparently exert any influence on the incidence of disease; so it was abandoned. Disease Prevention and Control During the summer and early fall, properly reared young grouse present such pictures of health as to lull the inexperienced breeder into a false sense of security. Early accounts contain innumerable records of entire groups being wiped out at this time, usually by the ever mysterious "grouse disease." The prevention and control of the diseases known to af- fect grouse during this period are today, however, so well understood as to furnish little cause for anxiety, providing a few important precautions are followed. Each pen should be thoroughly cleaned witli disinfcclant or firegunned in the fall in prep- aration for the following season. All grass and debris under the pens should be removed or THE INCUBATOR-BROODER METHOD OF RAISING GROUSE 495 burned. During the rearing season, flies, which may carry ulcerative enteritis, can be dis- couraged by the prompt removal of waste feed and the covering of the ground under each pen with a thin coat of hydrated lime once every two weeks. The price of freedom from most diseases is the maintenance of clean conditions within as well as about each pen. All feeding and watering equipment should, therefore, be disinfected at least twice a week and any accumulation of droppings either on the wire floor or on the conifers used as shelter should be promptly removed. As a precautionary measure it is wise to continue limitations on visitors and to place a folded feed bag in a pan of disinfectant so that all who enter the rearing field may step on it. It is, of course, hazardous to allow poultry or those caring for it, access to the immediate vicinity of the rearing field. In spite of every precaution, the two most serious epizootic diseases known to afiFect grouse in captivity, ulcerative enteritis and blackhead, occasionally may appear at this time. Until Dr. P. P. Levine, working for the Investigation, demonstrated in 1933 that the former might be carried by flies, even the most exacting precautions failed to check its spread. In 1932, 96 out of 196 birds were lost during September from such an outbreak. Each day, after this disease appeared all pens were disinfected, all feed dishes boiled and all feeding done by attendants wearing rubber gloves. These were dipped in disinfectant after servicing each individual pen. Yet the disease spread. During a similar epizootic the following year, all birds occupying supposedly uninfected pens were moved to a hastily fenced rearing field a quarter of a mile away where carefully disinfected pens were placed 100 feet apart. Even though a new attendant, who kept scrupulously clear of the old rearing field, was assigned to care for these birds, many subsequently were infected and died of this highly contagious disease. In 1934, however, it was demonstrated at the Research Center that by fly-proofing each pen with cheesecloth or muslin at the outset of the epizootic and by carrying out all the precau- tions normally associated with the control of an extraordinarily infectious organism, the dis- ease usually could be confined to the unit in which it originated. To make certain of control, uninfected pens may likewise be screened, although this practice was discontinued as unnec- essary at the Research Center. The key to effective control of ulcerative enteritis* is the prompt recognition of its presence. The very swiftness of the onset provides one of the best indications. Infected birds seldom appear sick more than 24 hours before death occurs. The unexpected loss of several young grouse from the same pen in a single night without apparent cause should always be a dan- ger signal to be followed by the immediate screening of the pen against flies. At this time a prompt examination of dead birds by an experienced pathologist may not always reveal ulcerative enteritis, for the characteristic lesions in the small intestine often do not appear in excessive numbers until the epizootic is well under way. Dead birds should never be kept within the rearing field. The only other serious disease likely to be encountered is blackhead. Birds reared in cap- tivity on the ground are most apt to acquire it, although even on wire a few are mysteriously affected each year. In the latter situation, isolating the birds from the infected pen into groups of two or three is usually sufficient to check it from spreading'''. Grouse thus infected may appear to be sick for several days since its progress in birds of eight to twelve weeks of age is not usually as rapid as is that of ulcerative enteritis. * See Chapter X, p. 437 for description and full control measures. A See Chapter X, p. 438. 496 ARTIFICIAL PROPAGATION OF GROUSE Very occasionally when accessible moldy feed has been allowed to accumulate, a fungus disease, Aspergillosis, may be the cause of death. Properly guarded against, one seldom has much to fear from any of these diseases at this period. Yet more of the early attempts to raise grouse artificially ended in failure through these causes than from all other difficulties combined — and just when favorable results seemed assured. Care during Rearing Constant attention to small items is the price of success during this comparatively unevent- ful period. It is wise to check conditions in the rearing fields several times daily. Rats are apt to congregate here because of the presence of waste feed beneath the pens. Tunnel traps properly screened are usually effective. Occasionally cyanide gas may have to be introduced into holes and closed runways to secure effective control. There have been several instances at the Center of raccoons amputating the legs of young birds. The feet were caught and pulled through the wire floor of the pen. By placing a single-strand electric fence outside of and close to the top of the rearing field fence these animals usually can be excluded. Dogs, cats and visitors, if allowed about the rearing field, may frighten the birds, causing head injuries. Though such scalping may be severe, the wounds usually heal. More important is the likelihood that the balanced social relationship so important in a pen may be upset by the scarred birds becoming increasingly nervous until a permanent inferiority complex even- tually is acquired. Such birds should be liberated for they seldom make good breeders. Birds quickly learn to know their attendants and to be reassured by familiar sounds when frightened. The practice of occasionally whistling or talking while working in the vicinity may also be used to quiet them during the periods of fright. Tame birds make the best breeders and reduce maintenance difficulties to a ininiinum at other seasons of the year. Considerable fear of man may be induced in young birds selected for liberation by frequent catching and handling or by placing a wild bird in the pen just previous to release. The Period of Social Adjustment The first cool days of early fall may bring into sharp relief the ever-present problem of social adjustment. Wild broods are then starting their fall shuffle or "crazy flight." The young males, previously seldom more aggressive than their sisters, now begin to display and to attempt to dominate their fellow penmates. Although fighting for supremacy may begin as early as the tenth week, the chasing and bullying of less aggressive birds is now apt to reach •S\'^ PRACTICABILITY OF ARTIFICIAL GROUSE PRODUCTION 497 a critical stage completely upsetting the social balance. The weaker individuals, bullied into attempts to escape, may be forced to spend most of their time on pen roosts or in the shelter of the pines placed in the pens for this purpose. Failing to reach such haven, they are occa- sionally severely scalped either by flying against the wire or by being viciously head-picked. As soon as trouble begins, the coniferous cover in the pen should be increased. Low corner roosts to which birds attempting to escape may fly, are also effective. By dividing the pen into several sections set apart by 10-inch base-boards placed on edge, the territory of the aggressive individual may be limited considerably, thus also assisting in reducing chasing. Occasionally the situation may be solved by segregating especially vicious birds for several days or by placing them in other pens where their dominant attitude tends to be neutralized by the resultant difficulties of establishing themselves in a new territory already occupied by strangers. Exposed to the proper brooding and rearing conditions, little difficulty should be expe- rienced with feather picking or cannibalism until the fail moult when the juvenile tail feathers are lost and the adult rectrices first make their appearance. Encouraged by the nervousness characteristic of the season, these, as well as the upper tail coverts, are often so severely picked as to cause considerable temporary injury. The prompt removal of both the offending and the injured bird for a period of a few days has already been suggested. To leave them apart for a longer period is, however, to run the risk of increasing difficulties in adjusting them- selves to their companions upon their return. Young grouse, to be kept as breeding stock, may be overwintered* in their rearing pens, particularly if two of these can be placed end to end to provide adequate flying space. Less labor and chance of accident are involved, however, if the grouse are brought together in large groups. A separate structure has proven most satisfactory at the Center for this pur- pose. Birds may be concentrated in winter quarters at any time after the first heavy frosts of fall have reduced the danger of a serious outbreak of ulcerative enteritis. In fact, provided quarters are larger, less trouble during the adjustment period will be experienced if this is done. PRACTICABILITY OF ARTIFICIAL GROUSE PRODUCTION Few will read this chapter without realizing that there are still several major difficulties to be overcome before the average game breeder can produce grouse in substantial numbers at a price within the reach of most individuals interested in restocking their coverts. A little time, however, may well be spent in analyzing production possibilities and limitations. The science of artificial grouse propagation is still too immature to permit an adequate evaluation of eventual possibilities. Sufficient progress has been made, however, to indicate that grouse properly handled in captivity will more than reproduce their numbers year by year. In 1940, at the Research Center, 65 hand-raised breeders laid 851 eggs, from which 502 youngsters were hatched. Of these 224 were reared to maturity (September 1). All but 37 of the latter were placed in breeding pens the following spring. It should be recognized that it has not as yet been found possible to maintain this pace each year. The care of the breeders, the incubation of the eggs and the rearing of the young all represent specialized problems, the successful handling of which requires experts with initiative, enthusiasm for the work, a flair for details and what is colloquially known as "grouse sense". Given all these, one still encounters "breaks", the reasons for which are not * See p. 459 for detailed description of overwintering. 498 ARTIFICIAL PROPAGATION OF GROUSE always clear, that strongly influence production one way or the other. Nor should it not be forgotten that the results in terms of birds produced are usually better with small numbers than with mass ])roduction. The picture at the Research Center was further complicated because it was necessary each spring to use part of the available breeding stock for one oi; another of the several experi- ments then underway. If uninterrupted attention could be given to the production of grouse for liberation, it now seems possible to rear from two to five birds to eight weeks of age for every female placed in the breeding pens. Production Equations For the reasons just indicated, it has not yet been possible to base a grouse production schedule on the record established at the Center. Each season some progress has been made experimentally in reducing the effect of the items which limit production. With such a stub- born problem, however, 13 years are still insufficient to learn to recognize and control all of the limiting factors. But those who would breed grouse artificially are naturally anxious to secure some basis for determining a logical production schedule. For such, the best guide at present seems to be the results obtained from experiments that have proved most successful after repeated trial. Given the proper conditions and supervision, it would seem quite possible to equal or to exceed the production indices here set up. TABLE 71. INDICES OF PRODUCTION Factors Number of adults in breeding stock Number of breeding; females Number of CKKs per female Total number of e^K^ I*ercenta(,'e of fertile ep^s —fertility Total number of fertile epjcs Percentatfe of fertile etr^rs Imtdiod — hat^^luibility Total number of eKj:s hat^lie>^^^ SIGNIFICANCE OF ARTIFICIAL PROPAGATION 503 no danger of extermination over most of its range except at the fringes. There still exist many areas where grouse were once known but to which they are now strangers. Whatever the cause of their disappearance may have been, it is always possible that the situation may have righted itself. Environments change with time. Such limiting influences as over-shoot- ing and destruction of the habitat through fire or grazing may be modified through conserva- tion education and agricultural adjustment to produce, once again, conditions favorable for grouse. The species is in reality much more sedentary than is generally realized. For this reason, unless a habitat from which grouse have been eliminated is adjacent to well populated coverts, the return of this game bird is apt to be a matter of many years. Where such coverts are isolated, successful reintroduction is dependent upon the liberation of new stock. This must either be trapped in the wild or reared artificially. As has been pointed out, few wild-trapped birds are available for purchase and those only at prices ranging from $10 to $25 or more a pair. However, there is seldom a valid reason for not trapping grouse from .sectif)ns where they are abundant but little hunted to be used for restocking depleted coverts elsewhere. The controlling factor here is the reluctance of most sportsmen to seeing any birds trapped from their localities. If a strain of grouse could be profhiced in captivity which could be raised in substantial numbers at a reasonable cost, and which was sufficiently adaptable to survive and breed, once liberated, this problem might be solved. This was one of our most important objectives. But the biological limitations of the bird, as already discussed, seem to be such that there is little likelihood that they can be modified except by an exhaustive program of selective breed- ing encompassing many years. For all practical purposes, therefore, one must conclude that, for the present, at least, the raising of grouse in captivity for restocking purposes offers little promise of producing sufficient birds at a cost reasonable enough to make it a prac- tical proposition. But on the bniad front there is much more to be gained than this end alone. Some of these considerations are so little recognized that they might well be listed : 1. Reasonable assurance that grouse can be maintained in captivity if tiiey are threatened with extermination in their natural environment. 2. Recognition of some of the less ohxious life hislor\ facts through association with grouse in captivity. 3. Opportunity for .studying the liiids piiNsiological and psychological reactions to the environment in which it is placed. 4. AvailahilitN of a much needed supph of experimental grouse uith which to carry on studies of the fuiidamciilal reactions of the bird to various en\ironmeiits. ^/?a«- '^Vf ^-^ 504 ARTIFICIAL PROPAGATION OF GROUSE There is. for instance, some indication that there exists an important relationship hetween the weijiht of crouse in the early winter and the livahilitv of the chicks hatched the following sprinp. Such a relationship may perhaps he hest measured hy a careful phvsiolojrical study where all factors can be controlled. For such a studv. captivity- raised birds must be utilized, for no method is known whcrcliN wild-trapped birds can be handled in captivity without producing serious physiological disturbances. Likewise, though one may determine grouse food preferences and composition from field studies and chemical analyses, it is not yet possible to translate these in terms of the nutritive value derived except by feeding various items to captive grouse and measur- ing the results. GROUSE LIBERATIONS The prime reason behind most efforts to raise wild game in captivity is the thought that it may some day lead to the production of a brood stock for liberation. The "increase in lib- eration" idea at times has worked well with some species such as beaver, deer, pheasant, quail and Hungarian partridge, though only where the habitat was suitable. Here, when the natural brood is short, artificial restocking may give just the boost necessary to speed recovery. The urge to produce grouse for this purpose was the spark that kindled most of the early grouse raising experiments. Onlv in the last decade have a few sportsmen, administrators and research men realized the added advisability of producing the bird artificially for study pur- poses. In reality too few grouse, either trapped in the wild or raised by man. have been liberated in depicted coverts to date to test adcquatelv the effectiveness of increasing the seed stock by this method. True, one may remember that 19 survivors of the liberation on .Anticosti Island in 1911 became the nucleus of an occasionally plentiful supply of grouse today; that, of five marked, hand-raised birds liberated in the fall of 1931 on a part of New York State's Con- necticut Hill grouse studv area, two were killed by ])redators. two survived the winter and one vanished. Most of the earlv introductions, however, disappeared or were absorbed by the res- ident population so that no real record of results were ever obtained. Transportation for Liberation No difficulty has ever been encountered in transporting either adult or young grouse raised in captivitv. In the early days of the Investigation the birds were carried five to a pillowcase on the back seat of a car. Later, shipping crates with wire bottoms and individual compart- ments for each bird were constructed. Most recently up to ten adults have been placed in standard pheasant shipping crates for transportation by express. Whatever the method, it is best to exclude all save a little light from the sbip]iing compart- ment to discourage feather picking. Burlap may be used to cover anv portion of the cage made of wire except the floor. If the birds are to be en route less than a day. they will feed or drink very little, so it is not necessary to make provision for either. Grouse from Nova Scotia and MaiiitDba |)laced in the comparlmented crates described above and given beechnuts or grain and green food, such as cabbage or halved apjilcs. were received in perfect shape at the Calskill Experimental Station. Water was provided in a small tin cu)). Tliis. though securely fastened to one cor- ner, should be removable to facilitate cleaning. Where such distances are involved, it is best to shi]) the birds in a wire-botlomcd crate to aMiid fiiuling the floor and soiling the feathers. Vi'herc mcire than one bird to a compart- GROUSE LIBERATIONS 505 ment is shipped, one encounters the possible danger of spreading disease and parasites, par- ticularly if the floor becomes dirty. Then, too, water spills through the wire floor, thus elim- inating further fouling. At present, most birds for liberation are obtained in Canada. Importation permits must be procured from the Federal Fish and Wildlife Service for those brought across the border WHITE OR ORANGE-COLORED CHICKEN FEATHERS, WIRED FAST TO THE TAIL FEATH- ERS OF THE GROliSE BEFORE LIBERATION. AID IN CHECKING ON SUBSEQUENT MOVEMENTS (rurdirifr Bump LIBERATED GROUSE SOON ADAPT THEMSELVES TO WILD CONDITIONS ALTHOUGH SOME NEVER REGAIN THEIR FEAR OF MAN SO CHARACTER- ISTIC OF THEIR WILD COUSINS into the United States. To avoid unnecessary delays it is wise to forward these permits either to the shipper or to the point of importation. Method of Liberation Although there is a thrill in the sight, no game bird should be encouraged to fly from the point of liberation. If possilile. the container should be placed where food and shelter are abundant and the birds allowed to wander out at leisure. Even with hand-raised birds it is not at all necessary to place feed in tlic innnediatc \icinitv. fi)r once in the covert they seem instinctively to sample buds, leaves and insects. A few released grouse may linger in the vicinity for up to a week, particularly if the feed to which they are accustomed is placed close by. There is no evidence, however, that such artificial feeding tends to encourage any 506 ARTIFICIAL PROPAGATION OF GROUSE large number of birds to settle in the immediate locality. Once liberated, most grouse will wander until they find suitable sections of the covert not currently occupied by others and then set up their own territory. They seldom seem to wan- der far for the rest of their lives. A number of birds thus liberated and later weighed several times were found to have lost from ten to 50 grams before adapting themselves to wild foods. Providing they are in proper condition, however, this loss is seldom serious. Care must be used to make certain that hand-raised birds can fly well. All broken primaries should be pulled out to allow for their prompt replacement. Marking for Identification It is best to release only banded birds unless there are no native grouse present. Size #9 aluminum leg bands are seldom lost if placed on birds over six weeks of age. Aluminum wing bands carefully attached to the inner webb in the bend where the humerus, the radius and the ulna meet, are also quite satisfactory when subsequent sight records of occurrence are not required. Where the birds are to be followed, tail marking* with brightly colored chicken feathers is also recommended as a distinct aid to future identification. But they are, of course, lost when the old rectrices are molted, not later than the following Julv or August. What Determines Survival Assuming that grouse are placed in suitable habitats, certain other considerations exercise a strong, though often little recognized, influence upon their subsequent survival. Notable among these are the source and experience of the birds, their age and their physical condition upon liberation. Success in dodging enemies, in finding food and shelter, in nesting and in raising a brood depends largely upon these points. Of course the inherent adaptability of the bird to new surroundings is also important. Source and Experience The first of these points, the source and experience of birds destined for liberation, is often difficult to control. Few birds ar*^ for sale and there are many who would buy them. It is almost axiomatic that the success of any colonizing effort depends in part upon the degree of similarity between the new habitat and the one from which the birds were taken. Thou- sands of dollars have been wasted in introducing southern quail into northern areas to which (hey were little adapted. Even though grouse may be a more versatile species in this re- spect, the chances of survival in a new habitat arc closclv tied in with tlicir ]irovious expe- riences. It is for this reason that hand-raised adults set.n to encounter consideraiil<' difficulty in making good wlicii lilicrati-d. Once they have grown up in association with man. many * Sec A|iprniUx. p. 717. GROUSE LIBERATIONS 507 seem not to acquire a strong fear of him, even after some time spent in their native coverts. Breeding birds escaping from the Research Center are occasionally picked up in the yards or gardens of the surrounding farms. One such bird traveled three miles during a period of several weeks, only to be caught quietly feeding in a greenhouse. Another, liberated on the Hyde Park estate of President Franklin D. Roosevelt, was killed in Poughkeepsie six miles away. Of the marked birds subsequently contacted from among the 191 hand-raised grouse, not necessary for propagation experiments and therefore available for liberation, about one- quarter never established a fear of man, one-half would flush at distances of from 25 to 50 feet when approached and the remainder compared well in wildness to native birds on the liberation area. With wild-trapped stock the situation obviously is somewhat different. Unless they are kept for a month or more in constant contact with people, they usually remain wild both in cap- tivity and when released. Only adults are trapped for restocking, for the younger birds from six to ten weeks of age are usually quite difficult to catch. Where the birds are heavily hunted and therefore difficult to approach, as in the Northeast, the Investigation found it practicable to trap birds for banding only on the nest or in the late fall or winter*. For this reason re- stocking with grouse in the younger age classes has seldom been attempted. Age The tanieness of artificially-raised adults mitigates against their utili/ation for liberation. Young birds, however, are much more adaptable than are the adults. In restocking a covert, the age of the birds liberated often exercises a measurable effect on the results attained. Most game birds raised in captivity seem to adapt themselves best to a new environment at a rela- tively early age. Experience to date indicates tliis may also be true of young grouse. There are no published records to indicate that any number of young grouse six to eight weeks of age have ever been released. At the Center, in order to secure the proper records of survival to be utilized as one criterion in the selection of the following years' grouse breeders, all young birds, except escapees, have been carried through to October before those to be liberated were chosen. Though few records are available to support the theory, it was observed that those birds accidentally liberated from the rearing pens in August usually went native quickly. Few of these birds were subsequently recaptured. Some were recognized months later by their col- ored leg bands. The indications are, then, that the best age at which to liberate captivity-raised grouse, is when they are eight to ten weeks old. Older birds are more likely to seek the habitations of man unless they are liberated at a considerable distance therefrom. Physical Conditions Another common cause for the failure of liberated birds to survive is poor physical condi- tion at the time of release. In 1931 the Investigation received a shipment of .34 Canadian birds, only 18 of which were in fit shape for liberation. That this situation is not unusual is indicated by many similar reports such as one from Martha's Vineyard where many of the birds received were described as "sick if not dead." Those released were never heard of again. Or. the other hand, two shipments from Nova Scotia arrived in excellent shape. * See Appendix, p. 716. 508 ARTIFICIAL PROPAGATIO\ OF GROVSE Distance, unsatisfactory shipping crates inviting injury and disease, and delays in transit sometimes due to importation difficulties all play their part. The condition under which the birds are held prior to shipment is also a factor. Ihiless all these considerations are prop- erly met, survival may be markedly lessened from the start. Ability of Liberated Birds to Survive In Chapter I liable 2, p. 18) one finds a record of some 2,300 grouse that have been lib- erated over the years, largely by sportsmen. These for the most part were wild-trapped birds. Yet a glance indicates that the number of valid returns covering the results obtained from this restocking effort is discouragingly small. In part this follows from the general failure to mark liberated birds in order to insure their subsequent identification. Likewise, only a few releases have ever been intensively followed to determine survival. Grouse, like pheasants and quail, even though plainly marked and faithfully followed are most difficult to find again following liberation. The only other basis on which results may be judged are the records of introduction that succeeded or were failures. The outstanding record of achievement already mentioned is the liberation of grouse on Anticosti Island in the Gulf of St. Lawrence. Most other releases have been into coverts where native birds were also present, thus making the attendant degree of success difficult to determine. Liberations are also being made and carefully followed in Ohio and Missouri but are not yet old enough to produce indicative results. Perhaps the most intensively controlled releases to date have been those made by the In- vestigation. Over the past 13 years, 191 hand-reared and 76 wild-trapped grouse have been liberated. These were made on areas where regular field surveys were being carried on by trained crews. In addition to banding, some of the birds were marked with brilliantly dyed chicken feathers wired or cemented to the tail feathers to facilitate subsequent identification. With the exception of 13 reared by Dr. Allen, the hand-raised birds came from the Re- search Center. Unavoidably, they were chosen from the surplus that remained after the breeding stock and birds for various experiments had been selected. Therefore, they might not be expected to do as well as birds turned out in the pink of condition. Nevertheless, of 84 released on the Pharsalia Game Refuge in 1938, 40 were subsequently contacted. One of these was four miles from the point of liberation. A marked female was found with a brood of seven chicks 78 days after release. The remains of 17 were found, though in most cases the cause of death was difficult to determine. In December 1932, on the same refuge, 29 marked, wild-trapped birds were released. At least 12 were still resident there the following April. In fact, four females nested and reared broods of young. Both of the above records are conservative as it is probable that some were not recognized because of the loss of identifying feathers. The latter birds wandered widely, one traveling three and a half miles in seven days while another was found six miles from the point of liberation a few months later. Others undouliledh siir\i\e(l Imt. liaviiiL' jnur- neyed off the survey area, were not contacted again. ^: ^ '*;^^.yM,.-f. . GROUSE LIBERATIONS 509 Already mentioned are the five hand-reared and -hve wild-trapped birds released in the fall of 1931. Two from each group were later found dead. Three of the wild and two of the hand-reared grouse survived at least until the following July. One was unaccounted for. One hand-raised grouse released on the Pharsalia Refuge nested the second year follow- ing liberation. This is worthy of note as the only record of this occurrence. Of the ten additional releases, all of which were carefully followed, the results were not radically different from those described above. In summary, of the 276 grouse liberated by the Investigation, 9.1 per cent were positively known to be alive a month or more after release. Though additional birds unquestionably survived, it is informative here to remember that Stoddard' ' found the hunter return from 2,516 imported Mexican quail, 680 native trapped and 211 hand-reared bobwhites to be but 6.3 per cent. Likewise, of 1,794 young male pheasants banded and released on New York State Landowner-Sportsman controlled hunting areas in 1941, only 362, or 20.1 per cent, were brought to bag during the subsequent hunting season. In addition to the above records of liberations made by the Investigation, there is a report, previously mentioned (Chapter 1. p. 191, of a release of ten males and seven females on Protection Island, Oregon, in 1939. The birds were followed closely and, although one produced a brood the year following liberation, they gradually decreased until but one re- mained in 1942. Hawks and owls apparently preyed heavily upon them and, while they be- came quite wary, they remain, ". . . too trusting of mankind . . .", and '". . . have not been able to withstand conditions on the island with success." In view of the above, there seems little room for sound conclusions based on present rec- ords of survival of liberated birds. We know that some birds will survive and a few, at least, will nest and raise broods. But it is yet to be determined whether or not hand-raised grouse, liberated into favorable habitats, can establish themselves or boost materially the num- bers of a depleted resident population. The general similarity of the results obtained from the release of hand-reared versus wild- trapped birds is interesting but not especially significant because of the small numbers in- volved. This is also true of the relationship between grouse, quail and pheasant contact rec- ords following liberation. Additional stocking then of first-class birds, particularly of the younger age groups, as previously suggested, on suitable areas must be carefully checked before one can adequately evaluate the possibilities of establishing or increasing grouse populations by restocking either with wild or with hand-raised birds. This is still a problem challenging to both wildlife manager and the grouse-conscious sportsman. \}i. CHAPTER XII PRODUCTIVITY OF GROUSE POPULATIONS By Robert W. Darrow POPULATION CHARACTERISTICS Composition — According to Age — Sex Ratios — Grouse Densities — Nest Densities — Brood Densities — Adult Densities — Sati RATION Point — Rate of Spread — Car- rying Capacity PRODUCTIVITY OF POPULATIONS Breeding Success — Survival During E\cn Life Period — Nest Period — Brood Period — Adult Period — Net Productivity — Life Equations THE BIOLOGICAL BALANCE SHEET & SUMMARY In the same way that individual grouse exhii>it certain peculiarities of l)eha\icir sn also do grouse populations have characteristic reactions to various influences and circumstances, (p. 513). Grouse productivity is the result of a continual tug-of-war between the forces tending to increase the population and those tending to rcdiici' it. I p. 551). The factors of increase include primarily the various components of the reproductive poten- tial of the species. Factors such as shelter, food and weather fall in this category only insofar as they are favorable. I p. 551 I. Decimating factors include predalion, disease, hunting and accidents, (p. 551 I. When one limiting factor (such as predation) is eliminated, others tend to take its ]>lace. ( p. 552 ) . It is jjrobably the cumulative effect of a number of small factors becoming active at the same time, rather than any one of them alone, that most often causes marked variations in grouse populations. ( p. 5.51 ) . Relative productivity from year to year tends to vary inversely with the density of the breed- ing population, (p. 540). When a fall grouse population numbers at least twice the corresponding breeding population it may be considered to represent a good crop. (p. 539). 512 I'liODUCriVITY Of GROUSE I'OI'VLATIONS The sex ratio lias aMnagcd nearly even dining liie Jnvesligatiuii luil a Iciulcncy has been noted for a regular seasonal variation to lake place, (p. 51 1). Average survival rates observed have been approximately 01 per cent for nests, 40 per cent for chicks and 46 per cent among adults on areas where no hunting occurred. (,p. 525, 527, 531). A considerable proportion of the hunting losses usually experienced appear to be deductible from the overwinter mortality which would otherwise take place. (,p. 58!vt. Individual coverts vary in their ability to support grouse populations over winter, depending basically on the quality of the cover. This concept has been termed carrying capacity. (p. 522). Densities of population tend to he higher in disconnected "woodlot" coverts than on wilder- ness range, (p. 520). Adults tend to avoid crowding above an average of one bird to four acres of grouse cover even in the best habitats, (p. 521). Grouse tend quite rapidly to move into a covert whose population has been depleted out of proportion to those surrounding it. (p. 522). Over a period of years productivity oscillates first to one side and then to the other of "dead center", (p. 546). Management may bring about a rise in the average population level but, once it has reached the improved plane, the tendency toward equilibrium will again assert itself, ip. 546). Real abundance, like real scarcity, is an abnormal situation, to correct which Nature is con- stantly striving for a balance, (p. 553 j. The potential productivity of most organisms is very high. As an entomologist has pointed out, a single pair of plant lice and its offspring could produce enough progeny in 12 years to cover completely the surface of the earth if all the various causes of mortality were removed. Grouse differ only in degree*. But in actuality unimpeded increase is seldom, if ever, approached. From its first appear- ance each species has faced constant conllict witli a more or less adverse environment. As a result it has become adapted to a distinct biological niche, entailing innumerable interrela- tionships with other species and influences. In this way its productivity has been controlled. As stated by Yapp'°°: "We may perhaps regard the organisms, both plants and animals, occupying any given habitat as woven into a complex but unstable web of life. The charaitcr of the web may change as new organisms a|)pear on the scene and old ones disajjpear duriii^i the ])hases of succession, but the web itself remains." The mechanism of regulation, however, is liigliiy complex iind is only bcfrinning to be understood. One of the early attempts to explain it was that of Malthus'" who came to the conclusion that populations are directly controlled by competition for food and that they always increase to the * Sec Chaplri Vni, p. 351. POPULATION CHARACTERISTICS 513 limit of susteiiancf. Iiis])ire(l by this |)rinciplp. Darwin liased on it his theory of natural selection''. More recent study, however, has shown thai, aside from a few exceptions such as deer which, in the absence of predators, have overbrowsed their winter range in certain locali- ties, lack of food is seldom a limiting factor. As pointed out by McAtee'™, only rarely is the solution so simple as Malthus supposed. Rather a host of interdependencies, both direct and indirect, are involved. Outstanding among them are those associated with the factors of shelter, food, weather, and predation. Less obvious are such influences as mating success, fertility and physiological resistance. Their combined interaction tends, on a long-term basis, to maintain the numbers of wildlife species in a state of equilibrium compatible with the biological niches to which they are adapted. In preceding chapters the various factors affecting grouse abundance have been considered individually. It is proposed to discuss here their combined effect on populations of this game bird as observed during this study. POPULATION CHARACTERISTICS As discussed in the chapter dealing with general habits, individual grouse exhibit certain peculiarities of adaptation and behavior. In the same way grouse po])ulations also have a typical pattern of composition as well as characteristic reactions to various influences and cir- cumstances. A knowledge of these is of the utmost importance to the game manager because in many respects they govern the extent to which management practices may be effective. Composition A fundamental factor in determining the productivity of any species is the age- and sex- composition of the population. This is particularly true for those which, unlike the grouse, do not breed their first year, but it is still of potential importance with these birds. According to Age In New York the great majority of grouse chicks hatch during the last week of May or the first week of June. These young birds have been considered adult by September and are cap- able of breeding the following spring. Therefore, populations of this species are made up entirely of mature individuals except during the summer. At hatching time the chicks have always considerably outnumbered the adults, the actual degree varying with breeding success and nest survival. On Connecticut Hill they have averaged 2.6 times as many, while on the Adirondack area the proportion has been 3.1:1, reflecting the usually lower nest mortality on this type of range. One should not assume, how- ever, that a degree of nesting failure sufficient to cause a reversal of this situation may not have occurred elsewhere or may not be experienced on the study areas at some future time. ' As the brood period progresses the preponderance of chicks diminishes until by the end of August the young birds usually but little exceed the adults in number. On Connecticut Hill the proportion* at this time has averaged 1.0:1, ranging from 1.6:1 in a year of high produc- tivity (1934) to 0.4:1 following a season of brood failure (1933). On the northern tract the average was 1.4:1 and the extremes 3.0:1 in 1936 and 0.4:1 in 1935. Data have not been sufficient to permit an analysis of age groups above the one-year level, although there is considerable evidence that among wild populations few grouse live more than three years'''. * Figures represent number of chicks for each adult, A See Chapter VUI, p. 360. 514 PRODUCTIVITY OF CROl SF. POPl L4TI0NS Sex Ratios Among newly-hatched grouse the proportion of males to females has been very nearly equal. At the Research Center this has been true both for chicks derived from wild eggs and for hand-reared stock. During tiie course of the Investigation the sex of birds encountered in the field has been judged so far as possible. But, as many a grouse hunter knows, there is no one sure way to tell males from females on sueh occasions.* Nevertheless certain criteria are available, partic- ularly in the spring. At this season especially and, to a lesser degree at other times, male grouse, when flushed, tend to rise rapidly from the ground while females tend to flv low for some distance. The sex of drumming and nesting birds is, of course, obvious. During the sunnner, too, most females are readily identified by the presence of a brood or by their broody actions. Summer data, taking into consideration also drumming and nesting information gathered during the spring, afford a reasonably accurate estimate of sex ratios at the beginning of the fall period. Data for the Connecticut Hill area are shown in table 72. In compiling this table, birds recorded as "sex unknown" have been omitted. TABLE 72. SUMMER SEX RATIOS OF ADULT GROUSE AS INDICATED BY POPULATION ESTIMATES— CONNECTICUT HILL AREA* —AUGUST 31, 1930-1942 Number Number Total Per cent Per cent of males of females sexed male female 1930 19 36 55 34.5 65.5 1931 79 77 156 50.6 49.4 1932 99 105 204 48.5 51.5 1933 26 33 59 44.1 55.9 1934 105 107 212 49.5 50.5 1935 120 142 262 45.8 54.2 1936 100 121 221 45.2 54.8 1937 52 93 145 35.9 64.1 1938 72 115 187 38.5 61.5 1939 S6 154 210 26.7 73.3 1940 99 115 214 46.3 53.7 1941 63 112 175 36.0 64.0 1942 72 94 166 43.4 56.6 Total.... 962 1.304 2,266 42.5 57.5 * Figures apply to entire area censused each year and. therefore, in many cases exceed those for the portion of the area used in tracing population fluctuations. The average ratio of 42. .S males to 57.5 feimles indicates that the summer grouse popula- tion on the Connecticut Hill area during the years covered contained more females than males. At the same time ratios as low as 34 males to 66 females (approximately), and as high as 49 males to 51 females, indicate that the relationship is far from constant. On the other hand, surveys conducted on the smaller Adirondack area failed to indicate a consistent predominance of females in the summer population (table 73). Because spring surveys on the Adirondack area were primarily designed to provide breed- ing population estimates, detailed drumming and nesting information was not recorded. Therefore, aside from females with broods, it was necessary to judge largely by flight char- acteristics when the birds were flushed. The number of individuals was low and the results as noted are inconsistent. X^IIiellicr these data are of sufncient weight to invalidate the idea » See Chapter U, p. 39. POPULATION CHARACTERISTICS 515 that the number of females exceeds the number of males in a normal summer population is a question. In all probability they are not. TABLE 73. SUMMER SEX RATIOS OF ADULT GROUSE AS INDICATED BY POPULATION ESTIMATES— ADIRONDACK AREA —AUGUST 31. 1932-1912 Year Number Number Total Per cent Per cent of males of females sexed male female 1932 IR 14 32 56.3 43.7 1933 13 10 23 56.5 43.5 1934 15 13 28 53.6 46.4 1935 5 9 14 35.7 . 64.3 1936 4 9 13 30.8 69.2 1937 7 11 18 38.9 61.1 1938 9 8 17 52.9 47.1 1939 14 13 27 51.9 48.1 1940 9 13 22 40.9 59.1 1941 9 II 20 45.0 55.0 1942 . . 12 10 22 54.4 45.6 Tntul 115 121 236 49.7 51.3 Supplemental data on this subject are available from the records of jrrouse that have been shot in connection with disease studies, population control and other sjiecial [iroblems. There is reason to believe, however, that much of these data is subject to errors which tend to accumulate rather than to compensate for each other. Thus, the use of a cylinder bore gun by several collectors over a period of years may have had more effect upon the se.\ ratio of the birds shot than did any variations which may have actually existed in the wild. Giving a wide shot pattern at close range, but with little capacity to kill at longer distances, its use resulted in a tendency to take the more closely lying females with somewhat more regularity than males rising at a greater distance. Therefore, instances in which the entire grouse popu- lation, or nearly so, of a covert was taken are a better basis from which to draw conclusions. Tn table Tt are summarized data representing the collection of practically the entire grouse population on a tract comjirising three comparlniciils totalling 1 ..S2(i acres adjacent to the Connecticut Hill sludv area between 1934 and 1936, one coniparlmeiit being covered each year. TABLE 71. Si:\ RATIOS OF GROUSE COLLECTED Dl RING NKARLY COMPLET ELIMINATION OF POPULVTION ON TRACT \DJ\CENT TO CONNECTICUT HILL— OCTOBER-MARCH, 193V-1936 Year Number of males Number of females Total* Per cent male Per cent female 19S4 1935 1936. ... IS 34 20 9 23 18 24 57 38 62.5 59.6 52.6 37.5 40.4 47.4 Total . . 69 SO 119 58.0 42.0 * Dnes not include birds so badly ahot'Up that sex determination was impossible. To the extent tiiat the tract on which the collections were made is representative of Con- necticut Hill as a whole, one may assume from these data that males exceeded females in numbers during the fall and winter on the basic sludv area also, but that there seemed to be a trend during the years involved toward a better balanced ratio. 516 PRODUCTIVITY OF GROUSE POPULATIONS It would thus appear that the sex ratio among adults at the end of the summer, in which females predominated, shifted to one having a majority of males between that time and early winter. Further, the fact that there has been no indication of hens suffering heavier decima- tion during the fall than cocks suggests that males outnumbered females among the maturing birds of the year. Analysis of collections of young grouse from .August through December since 1938* further indicates this possibility (table 7.5). It is recognized that these data are subject to sampling errors through selective shooting as noted above, although to a lesser degree because similarity in the habits of birds of this age is probably greater than among older birds. Never- theless the differences are sufficientlv consistent to be noteworlhv and statistically sound as demonstrated by analysis of variance. But the reasons for unbalanced sex ratios among matur- ing grouse are still obscure. •|\HLI'; 7.-). SE.\ It.vnOS OF IMMATURE GROUSE— STATEWIDE COLLECTIONS- AUGUST 1 THROUGH DECEMBER 31—1938-1941 Year Number of males 24 52 38 36 Number of females 19 43 32 30 Total Per cent mate Per cent female 1938 .. . 1939 .... 1940. ... 1941 ... 43 93 70 66 SS.8 55.7 55.3 55.5 44.2 44.3 44.7 44.5 Total ISO 124 274 35.7 44.3 If the trend so far suggested is real, then some influence must bring about another shift in the sex ratio from more males in winter to more females in summer. Such a factor seems to be a differential mortality in spring through which the cocks suffer more heavily. Of the 477 dead grouse (which were sexable) found on the Connecticut Hill area throughout the Inves- tigation and judged to have died from January through May. 59. .5 per cent were males. That this is quite likely associated with the breeding behavior of the cocks, especially drumming, is indicated by the fact that their mortality cui vc reaclii-^ it.- |ieak in \iiril while that of their mates drops off following a high in March". These data, therefore, indicate the existence of a basic tenileni\ fur llic sex ratio of wild grouse populations to become unbalanced in the spring through a disproportionate mortality among males, following which the balance is swung the other way by a jireponderance of this sex among the maturing young in the fall. Normally the ratio is most nearly even at some time in late winter or early s()ring. . It is likely, however, that this fundamental pattern is often modified by various circum- stances. For example, in a season of failure in the chick crop, the mitnber of maturing males might not be sufTicient to offset the summer surplus of females. Then the ultimate effect would be an augmented proportion of breeding females the following spring lending to compensate for the previous year's low priiduitivil\ . On the other hand, the greater the proportion of maturing young in tin- f;ill. ;in(l bcjuc llic number uf <'\lia males added, the more the resultant ratio would favor that sex. That such changes arc often associated with nuctualions in produclix il\ llicrc ran be little doubt, but it has been impossible from the data at hand to determine just how strong the * Thr l>iir*t n( Fabrirtis an a rriirrirtn of agr M.ifi imr knoun to the Invptiigatioii prinr to 1938, A Sri- itUcunaiiin nf ,A. .S16. POPULATION CHARACTERISTICS 517 relationship ma}' become. Tn any case, there seems small likelihood that, in years of increas- ing or high abundance, variations in sex ratio exert a controlling effect on grouse populations. Throughout the 13 years of the Investigation the proportion of breeding females on the areas under observation has at no time been seriously reduced. Nevertheless, one must bear in mind that a major period of grouse scarcity has not been experienced during this study. Grouse Densities Of interest also, to the sportsmen as well as the game manager, are the grouse population levels which may be attained in various coverts. It is often next to impossible to appraise adequately estimates of game abundance because they are not expressed in relation to some definite standard of measurement. What one observer may judge a plentiful supply of birds, another, with different experience, may consider only moderate. The data of the Investiga- tion are, therefore, expressed in terms of the acreage involved. The ratio of population to covert area is the density of population. In tracts of extensive forest, of which the Adirondack area is representative, the habitable range is usually the same as the total land surface. In disconnected coverts, which constitute most of the more heavily hunted grouse range in the Northeast, this is not true as only a part of the intervening fields is appreciably used by these birds. As explained in Chapter III. grouse cover in such localities has been considered to const^iute all woodland and lirush plus a 100- foot belt of the adjacent open land. This strip, on Connecticut Hill, amounts to about 29 per cent of all of this type present. Nest Densities As discussed in Chapter V. instances have often been observed where several grouse nested in quite close proximity to one another. But, in dealing with population behavior, coverts must be considered in their entirety. It has never been possible, however, to locate all the nests on the study areas. On the otlicr hand, it is believed tluil ncarlv all the females present in the spring have nested during most years*, although this jiruportion seems to have been somewhat low on Connecticut Hill in 1935 and 1936. Therefore, the number of this sex present during the breeding season represents a generally close approximation of nest density (table 76). It is apparent that nests are normally more numerous within llic disconnected tracts of grouse cover making up "woodlot" range than in extensive forest areas. This, of course, is also true of the average abundance of grouse. Nevertheless, wide variations are common in l)oth regions. With respect to individual coverts, the highest densities witnessed by the Investigation occurred on certain units of Connecticut Hill in 1935, even though the breeding population for the whole area was only the second highest observed. Just what levels were reached cannot be determined precisely, however, because no means has been devised for accurately judging the number of females failing to nest. The available evidence suggests that this proportion may have been above average in that year. In any event at least seven or more nests per 100 acres occurred on two units of 155 and 166 acres respectively, the number on the latter perhaps having been nine or ten as the density of breeding females was 11.5. On another unit of 131 acres, the corresponding figure was at least nine. During the same season several similar coverts held only three nests per 100 acres. At the other extreme a 230-acre « See Chapter VUI. p. 359. 518 PRODI CTIl IT) OF GROUSE POPULATIONS covert had only two nests in 1930, while in later years densities of less than two per 100 acres have not been unrommon on this area. TABLK 7(1. MIMBEH OK FKM\LKS PKH 100 ACHKS* I\ SPUING GROUSF POPULA.TION ON CONNECTICUT HILL AND ADIRONDACK AREAS— 1930-1912 Year Connecticut Hill AdirondackA 1930 2.1 3.4 5.2 6.5 4.1 5.5 3.6 2.6 3.0 3.7 3.9 2.6 3.2 1931 1932 2!i 1933 2.0 1934 1.2 1935 1.1 1936 1.7 1937 1.5 1938 1.6 1939 1940 1941 1042 Average 3.7 1.6 * In this table and throughout the following discussions of ilfnsity the data are given in terms of 100 acres of t:riMi9e cover. I'nits of this size are ren» 1939 l.:i 1940 1.1 1941 I) <) 1942 1.0 Average 1.7 1.1 * Only a portion of this area was worked in the summer of 1933. Supplemental evidence indicates that the brood density for the entire tract averaged only slightly lower titan in 1932. As Willi llic alimi(l;iiirc .if ilip species at all seasons, liro.ids have iisualK lieeii iiiarkedly less frequent in the forest hahilat of the northern area. Simihirlv their numbers have fluctuated less from year to year on that Irai I. On three occasions, however, their density on Connecticut ilill has dropped within liie ratine of thai on ihe .Adirondack area, hein;; jusl the same in 1010. Individual coverts have often exhibited higher, as well as l.>wer. densities than an) appear- ing in the averages for the area as a whole. Thu> as inan\ as 4.3 broods per 100 acres have been found on one tract of IRS acres while similar units have held onlv one in a number of instances and in 1930 none at all were observed on one of 231 acres. POPULATION CHARACTERISTICS 519 Brood density depends on the number of nests established, their mortality and the degree of renesting*. When nest losses are high, renesting may become quite important. This was apparently the case in several years when the number of family groups found during the sum- mer was greater than otherwise would have been possible in view of the breeding population and the nesting losses observed. On the other hand, there is at times a low abundance of broods in spite of high nest survival, as in 1935 when the data suggest that the degree of breeding failure may have been above average. Adult Densities The abundance of adult grouse is of especial significance at two times each year — the breeding season and the hunting season. In table 78 are presented the densities attained dur- ing the Investigation over the Connecticut Hill and Adirondack areas. TABLE 78. NUMBEH OF ADHIT GHOUSE PEM 100 VCHES ON CONNECTICUT HILL AND ADIRONDACK AllEAS— 1930-1912 Connecticut Hill* Adirondack Yoar Spring Fall Spring Fall 19.'50 4.6 6.5 10.2 12.1! 8..! 10.7 Ul 5.8 7.0 7.2 5.2 6.6 7.3 12.5 21.1 12.4 17.5 12.5 l.i.O 10.7 13.1 15.5 10.9 11.8 8.0 '4.9' 4.4 2.9 2.6 2.6 2.7 3.1 1931 7 4A 1932 8 6 1933 6 3 1934 1935 9.0 2 8 1936 7.8 19.37 7 2 1938 5.2 19.39 8 6 1940 1941 6 5 1942 7.4 Average 7.4 12.8 3.3 7.0 * Certain apparent discrcpanries between figures given in this table antl llinsc fur numerical populatinn eslini.nlrFt quoted in subsequent tables result fmni the (act that the total amount of grouse cover on the Connecticut Hill area in- creased during the course of the Investigation. A Represents January 1932 when first census on this area was made. Population estimates during the hunting season have not been made regularly. Sufficient work has been done at this season, however, to demonstrate that losses between September 1 and mid-October in general are minor. Therefore, data for the end of the brood period (August 31) when the young liirds become adult have been considered representative. Comparison of the two areas shows that grouse have been consistently more numerous over the woodlot range of Connecticut Hill. Beyond this, individual coverts have reached still higher levels. Thus in 193.5 the breeding population on a 166-acre unit was 22.3 birds per 100 acres while in other instances densities exceeding 15 birds have been recorded. Similarly in the fall a maximum of 38.8 on the same scale occurred over 187 acres at the end of the brood period in 1934. On another unit the figure was 31.5 in 1932 and a number of cases above 20 have been noted. It should be pointed out. however, that these very high densities have invariably been lowered through dispersion to surrounding less well slocked coverts, largely prior to the hunting s«ison. But such population equalization has had no observable effect on the average density for the area as a whole. As would be expected the minimum densities noted were associated with the generally low * The number of egps per cliilch and the number of cliicks hatching in successful nests have been quite constant. 520 PRODUCTIVITY OF GROUSE POPULATIONS grouse abundance of 1930. These were 1.8 and 2.7 for spring and fall respectively. On the Adirondack area the birds have occupied certain portions of the tract to a much greater degree than others*. For this reason the density over several hundred acres has been regularly somewhat higher than the average figures listed in the table. Likewise, throughout this type of range one will encounter pockets where grouse abundance closely approaches that of more diversified coverts. In the same way preference is often exhibited for particularly favorable situations in the smaller coverts of the woodlot type. Especially in the fall the hunter will frequently encounter concentrations within limited areas far exceeding anything attained over a range unit as a whole. Yet. even over larger tracts, abundance greater than observed during this study is indicated by accounts of others. Thus King"" has reported a bird to 1.8 acres in the fall (55.5 per 100 acres). In a letter to the authors Rowan'^ tells of a party of four hunters who, in the fall of 1941. killed more than 2.000 grouse in one week while operating from a single camp some 70 miles northwest of Edmonton, Alberta. But what is perhaps the most remarkable state- ment is that of Mershon^ who relates that in 1891 two hunters flushed 2,000 grouse in one day near Hemlock, Michigan. In addition to the two principal study areas a number of others have been censused at various times. These, although essentially similar in character to Connecticut Hill afford data from other sections of the State. The longest record is that of the Catskill area in southern Albany County where surveys were conducted in March of the years 1931-1941 except for 1936. Densities of adult grouse there averaged 6.7 per 100 acres and ranged from 2.5 in 1931 to 10.3 in 1933. Densities for the late winter season (February-March) for six years between 1931 and 1938 on the Pharsalia Game Refuge in Chenango County averaged 7.3 and ranged from 4.1 in 1931 to 11.1 in 1935. • In May 1934 a tract on Bull Hill in Tompkins County held 5.2 birds on the same scale. Supplemental information has also been obtained for range of more extensive character through the collaboration of technicians employed by the Resettlement Administration on two areas located on the fringe of the western Adirondacks and on one area in the western Catskills. Adult densities on these ranged from 1.8 to 4.7 in the spring and from 5.2 to 10.6 in the fall. Summarizing, the better grouse range of the woodlot type in the State may produce hunting season populations of upwards of 20 birds per 100 acres in individual coverts, although the average has been about 12. Yet even in years of generally high numbers, instances of scarcity are not uncommon. Abundance is usually lower over intermediate and wilderness range although local concentrations are often encountered. Most of the j)o|)uIation censuses in recent years outside of New York have been in Michigan and Minnesota. IIai>itat conditions in these regions are so different from those here that the results are not strictly comparable. Ncyprthelcss a number deserve mention. It should be noted that all of them eni])loyed the '"gridiron method" develojied by King"^" which is entirely difFerriil from the "((implcte strip" method u*ed in the New York lii\cstigation. Based on a survey of two square mile's of a township in the Pigeon Kiycr State Forest, Mich- • Sec Chnplcr Ul. p. 17.1. A Rowan, \('.. I'rnf<'>.or <•! Zoology, Unt%-eraily ol Alberta. POPULATION CHARACTERISTICS 521 igan, in 1933, Ruhl* estimated 3,444 grouse in the township, or nearly 15 birds per 100 acres (September). He states that the population on this area "during the 1933 season seemed to be high although the number was less than observed during the 1932 season." For the same area in September 1932 Fisher"' gives an estimated 191 grouse per section (sq. mi.) or 29.9 per 100 acres as well as 250 per section or 36.4 per 100 acres on a tract in Munuscong State Park (Mich.) in November 1934. In Minnesota, Trippensee^' gave 269 birds as the pre-hunting season (October) population of four sections of the Superior National Forest in 1934. or 10.5 birds per 100 acres, and 316 as the population of an area of the same size in the Chippewa National Forest, or a den- sity of 12.3. King"' states that the maximum breeding season population observed near Cliquot, Minne- sota, was a bird per four acres or 25 per 100 acres. The highest spring abundance recorded by Fisher"' was 15.4 per 100 acres in April 1933 on the Munuscong Park tract. Saturation Point From time immemorial hunters and others interested in wildlife have dreamed of game abundance far in excess of any it has been their fortune actually to encounter. Thus fanci- fully populated were the "happy hunting grounds" of the American Indian. Today's sports- men are no less given to wishful thinking. One of the prin(i|)al measures of the worth of a game management program is its effectiveness in increasing the density of such species as well as reducing the frequency and severity of periods of scarcity. Yet one must not expect the impossible. Regardless of how the habitat may be improved or the impact of decimating agencies relieved, populations will not increase above certain levels except sporadically. These levels and the ullimate limiting factors vary with the spe- cies. Deer, for instance, and other animals and birds which are highly gregarious may become very abundant before some environmental condition halts the advance. The rufled grouse, on the other hand, appears to exhibit an intolerance of crowding above an average of about one bird to four acres even in the best coverts. This has been termed the saturation point of the species and is a property of the bird itself apart from the quality of its environment. Evidence of this characteristic lies in the fact that, when greater densities occur, disper- sion of excess individuals to surrounding territory tends to take place immediately. The influ- ence, however, does not apply to birds of the year until after they have reached adulthood. Thus it seldom manifests itself except in the fall. While crowding almost invariably occurs within a brood group at the end of the summer, its numbers are usually accommodated within the same covert. But. when the covert a.^ a whole is overpopulated, wider movements ensue. During the Investigation, behavior of this kind has been observed on several occasions. Out- standing was the 187-acre covert which held 38.8 grouse per 100 acres as of August 31, 1931. Censuses later that fall showed its population to have dropped to 21.8 while those of adjacent units gained. Similarly in 1939 the density on a plot of 131 acres decreased from 29.5 to 20.6 while those of others nearby increased. That the birds will tolerate as many as 25 per 100 acres (approximately) was demon- strated when a portion of the Connecticut HUl area held a breeding population of 22.3 in 1935. King'^" has reported the same conclusion for an area in Minnesota. * Ruhl, H. D., personal letter to the authors July 2. 1936. 522 PRODUCTIVITY OF CROVSE POPULATIONS Rate of Spread Yet exceeding the saturation point is not the only cause of birds shifting from one covert to another. It is a basic axiom that "nature abhors a vacuum." Thus grouse spread from well-populated to unoccupied or poorly stocked territory unless for some reason the latter is definitely unsuitable. Because even adjacent coverts seldom follow the same trend in their population fluctuations such differences in relative abundance occur frequently. Accordingly, movements tending to equalize the distribution of the birds are continually taking place. But. because the net changes in population are usually small, it is difficult to follow them in detail. Although adjustments of this kind may occur at any time they are mainly associated with the fall and early spring seasons. In fact the terms "fall shuffle" and "spring shuffle" respec- tively have been used to describe them.* In general birds of the year are chiefly involved. The best indication obtained by the Investigation concerning the rate of spread into de- pleted habitats came in connection with a series of population control studies on land adjoining the Connecticut Hill study area. During the winter of 1933-34, 34 grouse were collected from coverts totalling 681 acres of grouse cover, reducing the number present to four birds by April. One month later 12 others had moved in and by the end of August further movements had increased the number of adults to 29, a gain of 25 in five months. The following winter, similar collecting was done in another covert of 572 acres. By March 1 there were 13 grouse left. During the ensuing month 13 more were taken but the population in early April was estimated at 17. Two additional birds were shot on April 7 and two others were lost to natural enemies during the spring reducing the number left to 13 by the end of May. Apparently none had moved in since March. By August 31 the estimated adult population had risen to 21. representing a total influx of at least 25 birds in six months. In 1935-36, the population of a third tract of 273 acres was reduced from 22 birds in Oc- tober to two in March. But in the interim, 24 birds had been taken. Thus four birds, in addition to those present at the start, must have moved in during the late fall and winter. Five more moved in by the middle of April, one more in May and one more in the summer. As the collecting carried on in these experiments was not begun until after the period of the "fall shuffle", these records demonstrate that apj)rociable shifts to restock depleted coverts may occur in March and early April and show that summer movements also take place. The consistency of the results on all three units indicates that such movements are customary and that any unoccupied territory in good grouse range will be quickly found by roving birds except possibly during times of widespread scarcity. The distance from which these inflowing birds had come is not known although in most instances they at least had to negotiate stretches of open land from a hundred yards to a quarter mile wide. That such distances are no bar- rier to grouse has been noted in the discussion of ni4. PRODUCTIVITY OF POPULATIONS 527 during this period by the occurrence of infertile eggs and embryos which do not survive in clutches the remainder of which hatch successfully. These have been of minor importance in the experience of the Investigation.* So also have been the eggs now and then filched by foxes and other predators. •ABLE 80. PROPORTION OF BROKEN-UP GROUSE NESTS OBSERVED DURING THE INVESTIGATION ATTRIBUTED TO VARIOUS CAUSE.S— 1930-1942 Cause Number Per cent Predation Man Fire... 551 40 13 12 3 2 88.8 6.4 2.1 1.9 Infertility Water O.S 0.3 Sub-total 621 27 100.0 Cause undeterminable Total 648* * A greater number of broken-op nesls were observed then were comparable for correlation with the number known to have hatrhr4 1935 1934 1937 1938 1939 1940 1941 I94Z YEAR FIGURE 43. GROUSE BROOD SURVIVAL RECORDED ON CONNECTICUT HILL AND ADIRONDACK AREAS —1930-1942 The primary causes of losses at this age, however, are still a major problem. Predation seems to be responsible for rouphlv half the mortality in averajie seasons. Few specific data have been obtained to account for the remaintier. As noted in Chapter VII (p. 316), a sub- stantial parallel has been observed between the early decline among grouse chicks hatched and reared artificially at the Research Center and that among wild broods. But even here the closest diagnosis which has been possible so far is the ambiguous term ''nutritional defi- ciency" which seems actually to refer to some physiological disturbance as sufficient food is always available to these birds. Also studies of the foods eaten by wild chicks have indi- cated a superabundance. Another influence is weather but its significance has not been de- termined, although there can be little doubt that the cloudburst in central New York in early July. 1935, directly increased the mortality of grouse chicks. Furthermore it seems most likely that the markedly lower survival experienced in some years may fundamentally be the result of some combination of these other factors rather than of predation. There is no evi- dence, however, that disease has been important. Among the relatively few chicks found dead on the study areas, no cause of death other than predation has been noU'd. althougii in 21.7 per cent of the cases no diagnosis at all could be made. Elsewhere fatalities resulting from highway and miscellaneous accidents have now and then been observed. It has been suggested that, as the number of chicks hatched |>cr unit of area decreases, the corresponding proportion maturing tends to increase. Ihc data, howcycr, do not support such a conclusion as shown in table 82, in which the records for the Connecticut Hill and Adiron- dack areas were grou])ed according to the relative alunidance of chicks at the beginning of the brood period. In fact on the northern tract, better survival was associated with the higher PRODICTIVITY OF POPULATIONS 529 densities. In both cases the years included in each group were well distributed throughout the period of study. TABLE 82. RELATIONSHIP OF DENSITY OF GROUSE CHICKS AT HATCHING TIME TO SURVIVAL DURING BROOD PERIOD ON CONNECTICUT HILL AND ADIRONDACK STUDY AREAS— 1930-1942 Connecticut HiU Adirondack Group* Average number of chicks tiatched Density per 100 acres Average number of chicks surviving Per cent survival Average number of chicks hatched Density per 100 acres Average number of chicks surviving Per cent survival I II 290 400 S48 12.1 16.6 22.8 108 199 37.2 37.3 36.3 70 96 8.6 11.8 25 38 35.7 39.6 Ill * Croup I — years having a density of chicks at hatching oo Connecticut Hill of under 15 per 100 acres, on the Adirondack area oi under 10. Croup II — years having densities between IS and 20. and over 10 reitprrtively. Croup III — years on Connecticut HiU with densities over 20. Nevertheless, the trend in the number <>f chicks left as of September 1 each year has fol- lowed in general that of the number hatched (figure 44). This was not true on Connecticut Hill in 1935, however, when a marked increase in the rate of mortality occurred as a result of abnormal weather conditions. The same season witnessed a very low survival on the north- ern area, too, although the number of chicks hatched was the lowest recorded. Survival was poor on both areas in 1933 also. On the other hand, if the years are grouped according to the density of the breeding pop- ulation— in which case wide ranges of chick abundance are included together — one finds an inverse relationship with respect to the percentage of brood survival (table 83). The degree to which this may represent cause and effect as well as the reasons which may be involved, however, are not apparent. One possibility for speculation is that the forces responsible for lowering the breeding population level may tend to result in an increase in the average stam- ina of the remaining stock. A similar condition among the progeny of these birds might then account for the better brood survival associated with them. & 530 PRODUCTIVirV OF GROISE POPILATIONS — — Number CHickft Hatched — Number Surviving •^ — Rorcent Survival CONNECTICUT HILL ADIRONDACK (J I .50 u 100 (T liJ ID 50 Z o le^O I9M I9SZ ieS9 ISSA ISSb I9S« 1937 l^^a 1939 \940 1941 tS-i-Z YEAR FIGURE 44. RELATIONSHIP OF NUMBER OF CI1K.K> HATCHED TO M MBER AND PERCENT SIRVIVING AT END OF SUMMER ON CONNECTICUT HILL AND ADIRONDACK AREAS — 1930-1942 PRODUCTIVITY OF POPULATIONS 531 TABLE 83. RELATIONSHIP OF DENSITY OF BREEDING POPULATIONS TO SURVIVAL DURING THE BROOD PERIOD ON CONNECTICUT HILL AND ADIRONDACK STUDY AREAS— 1930-1942 Connecticut Hill Adirondack Group* Average density of breeders per 100 acres .\verage number of chicks hatched Yearly variation in chicks hatched Per cent survivalA Average density of breeders per too acres .\verage number of chicks hatched Yearly variation in chicks hatched Per cent survival I 5.2 7.2 10.6 366 431 484 200-474 264-625 330-596 40.7 37.9 30.8 2.8 4.7 62 81 60-100 8.3-109 50.0 II Ill 38.3 * Group I — years having a breeding population density, on Connecticut Hill ol under 6 per 100 acres, on the Adirondack area of under 4. Group 11 — years having densities between 6 and 10, and over 4 respectively. Group HI — years on Connecticut Hill with densities over 10. A Analysis by chi-square shous that on Connecticut Hill Group 11 was very close to the total average while Groups i and 11! were signiBcantly different. Adult Period In appraising adult survival, September 1 each year has been taken as a starting point be- cause this is the time the voung of the year are considered to "come of age". By the same date the following year slightly less than half have been left as an average, most of the loss having taken place prior to the breeding season (table 84). On Connecticut Hill, however, a much higher survival followed the low fall populations of 1930 and 19.31. A similar rela- tionship was observed on the Adirondack area in 193.5-36. TABLE 8t. ADULT SURVIVAL RECORDED ON CONNECTICUT MILL AND ADIRONDACK STUDY AREAS^193(i-19H* Area Adult survivjil data Year 1930 1931 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 Aver- ajrc Initial September 161 276 465 274 420 300 311 273 334 394 276 .300 31S Connecticu t IliU Per cent survival: 88.2 85.7 81.5 71.1 60.9 42.4 72.6 59.5 61.0 47.4 56.7 48.0 39.5 37.0 54.2 57.2a 53.0 48.8 46.7 41.2 47.8 45.3 55.7 42.3 58.4 to September 49.8 Initial September population 60t 70 51 73 23 64 59 42 70 63 S3 57 Adirondack Per cent survival: to April! 66.7 53.3 51.4 37.1 47.1 56.9i 28.8 21.9 91 3 69.6 34.4 28.1 42.4 42.4 71.4 34.3 39.7 43.4 47.4 to September ... 42.1 *Year indicated is that of the beginning of each September 1 to August 31 period. Alti 1933 on the Adirondack area and in 1937 on Connecticut Hill the number of birds moving into the area during the summer e.\ceoded the iiilliiher k)Sl. tl^stimate made in Jtinuary 19.'i2 and therefore somewhat below September level. jSpring censuses were discontinued on this area after 1938. As has been discussed in Chapter VII, some 80 per cent of the losses during this period have been attributable to predation, the remainder resulting largely from accident and disease. Just what the relative importance of these minor causes has been, however, is difficult to judge because few of the dead grouse located during the survey have been found in a thoroughly fresh condition that would permit recognition of disease. On the other hand, field experi- ence indicates that accidents are infrequent and the collection of specimens for pathological examination has revealed a very low incidence of disease. At the same time one must re- 532 I'RODI'CTIVITY OF GROVSF. POPULATIONS member that the effectiveness of all these decimating agents is affected in varying degrees hy a number of factors such as cover quality, weather and buffer abundance. The basic data of llic Imcstigation are not influenced by the factor of hunting since this has been prohibited on the jHJinarv study areas. Neither has there been any evidence of starvation observed. The following table summarizes the analyses of the dead grouse picked up. Some of the specimens charged to predalion very likely died of other causes but the available evidence gives no reason to believe that the proportion of such instances has ever exceeded the dif- ference between the figures listed below and the 15-odd per cent estimated to cover such losses*. TABLK 83. PROPORTION OF DEAD GROUSE FOUND ON VARIOUS STUDY AREAS ATTRIBUTED TO VARIOUS CAUSES— 1930-1942 Connecticut Hill Adirondack Other areas Total Number Per cent Number Per cent Number Per cent Number Per cent Predation 446 17 20 92.3 3.5 4.2 16 100.0 125 100.0 587 17 20 94.1 2.7 Disease 3:2 Sub- total 483 450 100.0 16 14 100.0 125 86 100.0 624 550 100.0 Cause undeterminable. Total 933 30 211 1174 On the various study areas, causes other than predation were identified only on Connecti- cut Hill. The instances of accidental death are all attributable, so far as is known, to col- lisions with trees, fences and sometimes buildings. Also they have occurred chiefly during the fall and, in most cases, appear to have been associated with the so-called "crazy flight". Losses resulting from crippling by hunters have not been a factor on the areas from which this ma- terial was collected, although on heavily shot tracts they undoubtedly enter in'^. Incidental records from elsewhere over the State have also involved a number of other types of accident. Along improved highways birds are often struck by automobiles. Females have been found burned to death on their nests following forest fires. And. now and then, a grouse has been caught in a fox trap. The disease relationships observed are discussed in Chapter X. The number of adults surviving from year to year has been only slightly correlated with differences in the level of the preceding fall po|)ulation (figure 45). On the other hand, the number of adults lost each winter has borne a marked relationship to fall abundance. As discussed elsewhere''', other conditions being similar, the more numerous grouse are the more frequcntb they will be encountered and killed by predators. The data for each year on the two main study areas are compared in figure IC)*. In both cases, as the fall population has risen, so also has the mortality. Yet the proportion lost has iiccn less related (figure 17)'. Although the trend has been similar, the degree of change has b<-en less and exceptions more frequent. The correlation has been stronger on the Adirdiidack area than on Coiuiccticut Hill. At the same time, when iIk- data were analyzed on the basis of average values after the • Sec Cho|.l.r Vn, [.. 319. A Sec Chaplrt IX. !>. .176. t Src Chopler VII. p. .121. t BertUBC ol the wide tlifTrrence in the numerical popuUtiona involved (»n lhi>«r two arean the tinia covered been rediirrit to percentage deviation from the mean. thia graph have PRODUCTIVITY OF POPULATIONS 533 Foil Populafion Sunvival +o Followinq Foi 500 400 U) 30O a CD U. Ozoo Z loo CONNECTICUT HILL 500 400 300 ZOO loo I I I ' ' I «/) loo a. ID o q: u IS D Z 50 ADIRONDACK loo bo I930-31 Sl-32 3Z-33 33-34 3435 3&36 3037 31 38 38-39 39-40 40-*l 41-42 YEAR FIGURE 45. RELATIONSHIP OF ADULT SURVIVAL TO FALL POPULATION LEVEL ON CONNECTICUT HILL AND ADIRONDACK AREAS — 1930-1942 records had been grouped according to the density at the beginning of the period each year, further evidence of a fundamental relationship of this kind was revealed (table 86). Again it was more pronounced on the northern area. It is on this basic tendency for the proportion of mortality to vary with the fall population level that the concept of carrying capacity* depends. That the figures have in many instances * See discussion of Carrying Capacity, p. 522. 534 PRODUCTIVITY OF GROUSE POPULATIONS z ^s < u z^ '" 02 <^ ^^ I- z u '^ a. UJ is Z5 50 - CONNECTICUT HILL _!_ ADIRONDACK T5 - SO 2S o Z5 50 75 _l_ -L. -50 o -t-so -50 o +50 NUMBER LOST PERCENT DEVIATION FROM MEAN FIGURE 40. RELATIONSHIP OF FALL POPULATION LEVEL TO NUMBER OF ADULTS LOST ON CONNECTICUT HILL AND ADIRONDACK AREAS — 1930-31 TO 1941-42 25 Z < LI 75 zi ^o g a: l-i^ il Q. I- O o< °-> _lQ 100 D 2 CONNECTICUT HILL 300 ZOO loo 10 o u D Z I I ADIRONDACK too So 1950-31 3I-3Z 32r33 33-3-i 34-35 3536 3f the dead grouse picked up on the survev areas it has been possible to estimate quite closely the time of di'alh. Tliix- (lata arc jilottcd in figure 50*. No important variations between years have been noted. * On arrna nprn to hunlinK fl minor htimp in thr fur%-e during liir open drnson wonlH be expected. no o z > a: D 50 V) CO _l D 5 30 I- z 20 til o cc 10 u a. I PRODUCTIVITY OF POPULATIONS Cor>n6crl-icu+ Hill Adirondack 537 n ^o I o "J u _L -L. _1_ CM 60 40 30 zo 10 l930-»t 31-32 3i33 33-34 3+36 3S3f brcrders a» in table 8.1. It is nolt'worlln that on Cunncctic iit lliil as tlic density of the spring; |)i)[)ulali(.)n increased so also did the proportion of the loss in productivity which look place during the breeding * On Cnnnprlicut Hill thi* rorrrlatinn in tlalittically signifiraDt wtiilr on tlir othrr aret it it ffomewhat Icm drBnjte. A Analysii of these by clii-squarc ■huwt that on Coniircticut Hill Croup H baa been very close to ibo total average, but the drviations of Groups 1 and HI arr hiichly significant. PRODUCTIVITY OF POPULATIONS 541 and nesting season. This occurred in spite of the fact that, as shown in table 83, the actual degree of brood mortality recorded also increased as the breeding population rose. On the Adirondack area, however, there was no valid difference*. Considering these observations in the light of the fact that the trend in the production ratio has been inverse to that in the number of breeders (table 88) certain inferences may be drawn. On the Connecticut Hill area, with its generally higher population level, losses prior to the brood period appear to have been the stronger influence with respect to the greater average reduction in productivity observed to be associated with higher densities among the breeders. On the Adirondack area, however, even though the net productivity declined as total losses increased with higher densities, the relative importance of the breeding aiid nest- ing season as compared with the brood period did not change materially. At the same time, because the percentage of brood mortality itself has been very similar from year to year on the two areas, the fact that it has represented a greater proportion of the total loss of produc- tivity on this area suggests that the higher nest survival usually experienced in the Adiron- dack region may have been largely responsible for the greater average production ratios recorded. Since the relationship of density to productivity may be of major importance in grouse management special efforts were made to evaluate it. After virtually eliminating the birds over 681 acres of coverts adjacent to the Connecticut Hill study area during the winter of 1933-34, the population unit was (ielibcrately reduced during each of the succeeding three winters well below what appeared to be its carrying capacity. The data for 1935, 1936 and 1937 are compared in talile 90. that for the first year not being comparable because of the rapid influx of birds from surrounding territory'^. I XRr.K 00. EFFECT OF HltKKniNCi I'OPHLVTION HKIMICTION ()\ PHODUCTIVITY AS OF si;i'ii:\ii5i';u i c.oNNEc.iif.rr mi.i, I'w.i-i'UT Controlled area Check area Year Number breeders Production ratio Number breeders Production ratio IQSS 1936 45 38 28 1.91 2.16 2.29 256 170 123 1.17 1 82 1937 2.21 Several aspects of this record are of considerable interest. Each year the production ratio was greater on the controlled unit than on the basic study area. Furthermore, it increased from season to season as the number of breeders was reduced lower and lower. Yet it in- creased on the check area also until in 1937 the differential between the two was negligible. This, too, was accompanied by a progressive decline in the spring population. In view of the fact that the two productivity rates came closer and closer together each year it is note- worthy that the actual density of breeding birds on the two units followed a similar pattern in spite of their both decreasing. Further analysis of the records reveals another significant fact. The season of greatest difference in relative increase, 1935, also witnessed heavy brood losses, apparently as a re- * Data were analyzed by clii-squaie. A See disrussion of Rale of Spread, p. 522. ?>V2 PRODUCTIVITY OF GROiSE POPULATIONS suit of the Julv flood. The average number of chicks ])er brood inaluriiig, however, was simi- lar on both plots. On the other hand the percentage of females rearing broods on the con- trolled unit was nearly double that on the unmanaged area (table 91). The same relation- ship prevailed the following two years, though less strong. These data indicate that, during the same year when environmental conditions were uniform over both units, the greater pro- ductivity associated with lower de:isities of the spring population was largely a result of more successful breeding and nesting rather than of greater survi\ al among the chicks. TABLE yi. EFFECT OF BREEDING PUI'LIATION REDUCTION ON PER CENT OF FEMALES REARING BROODS AND AVERAGE NLLMBER OF CHICKS PER BROOD MATURING— CONNECTICUT HILL— 1935-1937 Controlled area Check area Year Per cent of femnlps rearing broods .\v»Taj;e iHimbiT of chicks uiaturiug Per cent of females rearing broods .\verage number of chicks maturing 1935 69.5 47.6 68.7 2.8 4.7 3.7 38.9 40.2 56.7 2.0 1936 4.8 1937 4.2 At the same time, however, it has so happened, as shown in table 83, that survival among the chicks on the study areas over the full period of the Investigation has varied inversely with the density of the breeding population. Yet the extent to which this represents cause and effect is not clear. In any event, on Connecticut Hill, it apparently was the weaker in- fluence, on the average, in reducing the productivity of high spring populations, although this does not seem to have been the case on the Adirondack area. With respect to the relative roles of breeding failure and nest mortality, lack of suffici- ently precise data makes appraisal difficult*. On Connecticut Hill there seems little doubt that in 1933 both were high. In 1935 and 1936 the former seems to have been high and the latter low, while in 1940 the reverse was the case. In 1912, the other year of especially low productivity, both were average but brood mortality was high. On the .Adirondack area both have been consistently low. To sum up, it appears that in the better grouse coverts of i\ew York, supporting as they do higher densities, the increment of young birds little more than equals the number of breed- ers even in years of good productivity. In the more sparsely inhabited .Vtliroiulaik coverts, on the other hand, it has averaged nearly half again as great. Now and then, however, the forces limiting |)ni(hictivity result in virtual failure of this crop. Heyond this, the relative in- crease on a given area tends to vary inversely with the density of the breeding ])opulatiL I'OPl LATION DEGREASIM, PRIMARILY AS A RESULT OF HIGH OVERWINTER MORTALITY i74 Breeders infertile — deod qerros DECREASING POPULATION ! Hiq\o r)es\ loss 140 Br-««der3 FIGURE 55. AVKRACK I.IKE EQUATION OF CROl SE POI'l I.ATION DECREASING I'RIMAIill.Y AS A RESULT OF III(;M NKST MORTALITY 174 Breeders rD^^breediog loilure dead cx'srr^^ WonVei DECREASING POPULATION (High br-ood loss 125 BreedetrS FIGURE 56. AVERAGE LIFE EQUATION OF GROUSE POPULATION DECREASING PRIMARILY AS A RESULT OF HIGH RROOD ATORTAT.TTY 550 PRODUCTIVITY OF GROUSE POPULATIONS portion, as discussed earlier in this chapter (p. 538), observations indicate that about half, or 9 per cent, represents a loss which would occur anyway. Therefore, in the diagrams the hunter take is shown in full while 9 per cent has been deducted from the overwinter mor- tality as derived from the study area data. The data indicate that this degree of hunting is seldom a decisive factor. Even under the situations represented by figures 55 and 56 its elimination would still fail to prevent a net loss of breeding stock. On the other hand, as populations approach scarcity in an area the importance of hunting increases and may become limiting. Under such circumstances a low degree of productivity by the end of the brood period would be the cue for reducing hunting pressure. But while the general population level remains high even failure of the increment of young birds does not necessarily mean that shooting should be curtailed. For example, while the September ])roduction ratio on the Connecticut Hill area in 1933 was only 0.96, it followed the highest breeding population recorded and the actual fall density was greater than those of 1937 and 1940 when the production ratios of lower breeding populations were 2.21 and 1.50 respec- tively. There seems little doubt that in all three of these years the birds could have withstood average hunting without endangering the prospect of a good crop the following season. The possibility that a greater harvest might on occasion be permitted, withort impairing the breeding potential, depends on the size of the fall crop. The foregoing equation may be modified to indicate the latter by restricting the term AL (adult losses) to cover only the period from April to the beginning of the hunting season. But practical interpretation of estimates at this time depends on a knowledge of the portion constituting a surplus. Means of evaluating this point are discussed in the section of this report dealing with management*. THE BIOLOGICAI, BALANCE SHEET-^ In preceding chapters the observations of the Investigation with respect to the major fac- tors controlling grouse abundance have been discussed individually. In the present chapter the data relating to survival, productivity rates, and life equations have also been consider- ed. It seems logical at this time to undertake a brief summary of the host of interactions among the various competing influences involved. The situation may be likened to a picture puzzle composed of a great variety of pieces, some large, some small, with a few still hiding unrecognized in the box. The title of the picture is "The Grouse Crop." On the skill and accuracy with which the pieces can be fitted together depends the solution. But the game is made more difficult and interesting because Nature has provided an infinite variety of basic patterns, constituting grouse cover on which the pieces, like semi-transparent overlays, may be fitted together. For each such pat- tern the pieces combine a lilllc dilTcrcntlv so lliat. in reality, each one represents a puzzle in itself. Standing at one"s elbow in this game are many players, both amateur and professional, each convinced that the solution of the various puzzles requires a different arrangement of the pieces (such as shelter, food, predalion or disease). Guiding rules as to the soundness of conclusions regarding relationships must be developed through experience and study. It is the intent of this book to define and explain as manv of these rules as the a be affected depends on the severity and duration of the forces involved. Yet these are relative, constantly conditioned by the end- less variety of circumstances. For instance, when grouse are scarce, a hunting pressure thai may mean little to them when they are abundant, may be the drag that prevents their in- crease. On the other hand, excessive populations appear more susceptible to disease. 552 PRODUCTIVITY OF GROUSE POPULATIONS While there can be little doubt that predators are the chief executioners of grouse, except during the early brood period, this game bird can be considered little more than a delicacy or dessert to most of them. At the same time a host of other influences are responsible for fluctuations in the vulnerability of the birds. The effect of shelter quality has been noted as has also that of the density of the grouse population. Weather, too. can be fayorable or un- fayorable. Another highly important factor is the availability of buffers. Usually more nu- merous and easier to catch, they constitute the staple diet of predators as long as the supply lasts, even though grouse may also be plentiful. But let the buffers decrease substantially and the fox. weasel, great-horned owl and goshawk, harder put to keep from starving, make more frequent and more determined efforts to capture grouse. Thus an epizootic, decimat- ing the ranks of rabbits and mice, may indirectly exercise a strong influence on partridge abundance even though predator populations remain essentially unchanged. That predators are more likely to catch birds slowed down by disease there can be little doubt. Though such cases do not appear to be as common as is generally supposed, the re- moval of afflicted birds from the coverts obviously is beneficial in inhibiting a further spread. In America, the usual restrictions on hunting are still so fixed as to discourage the taking of a large part of the crop when appreciable surpluses occur. Thus, during the more produc- tive years, disease- may actually be encouraged by leaving an over-abundance of birds in the field. To the extent that predation serves to whittle down this excess, it may help to maintain a goodly number of birds. So clearly is this situation recognized in Great Britain, where pred- ators are scarce, that gamekeepers each year spend considerable time in "shooting down" the population of Scotch grouse, left at the end of the hunting season, to a level where out- breaks of disease are unlikely and the carrying capacity of the moors is not exceeded. Furthermore, reduction of a decimating agent is no guarantee of a corresponding in- crease among the grouse. Compensatory adjustments soon tend to take its place. For ex- ample, unless applied to a large area, the effects of predator control are rapidly offset bv the influx nf individuals from surrounding territory. Similarly, as was observed on one study unit, if only certain species such as foxes and weasels are removed the depredations of others such as skunks and crows increase. On Valcour Island (N.Y.). where this relationship was also studied, after the elimination of all the usually more important predators, the propor- tion of nesting loss attributable to red squirrels increased. Even if some combination of circumstances should allow an excessive density of birds to build up in the fall, heavy overwinter mortality may usually be expected to ensue unless the majority of the surplus is taken by hunting. It seems probable, too. that the occurrence of high concentrations may often set the stage for serious outbreaks of disease, \ature is constantly trying to balance her books. One cannot examine this picture without becoming conscious of il> licnicndous complexity. There is little solace in it either for the sentimentalist who would stop all hunting or for the sportsman who thinks an abundance of birds can be assured through predator control, for the data clearly show that Nature's grouse budget can never be increased for long by either means. Certain factors, such as weather, accident and the productivity of the bird, are be- \ond human control. ()\er others, such as hunting, disease, predation. buffer abundance, food and shelter, the game manager may exercise varying degrees of regulation. Rut grouse populations are dynamic, never stable. Experience indicates that their fluctuations are not easily predictable, so that measures taken to allcyiatc scarcity or maintain abundance are of- ten too little and too late. THE BIOLOGICAL BALANCE SHEET 553 The prospect, however, is not all dark. The reproductive capacity of the grouse repre- sents an ever-present source of potential increase. By making poor coverts good and good coverts better Nature can be encouraged to set up a relatively high balance in which a dense population of grouse is the rule rather than the exception. Her efforts can be assisted by eliminating the hunting of grouse during periods of real scarcity and, it appears probable, by increasing the harvest of the crop when it is abundant. The conclusions and recommen- dations of the Investigation along these lines will be found in the following section of the book which deals with management. In summing up, one must realize that each grouse covert has its own balance sheet and that it is the sum of these, rather than the number of birds which may be present in one's own favorite hunting ground, that determines the size of the total grouse crop. Likewise the degree to which the average level of abundance may be maintained is dependent not so much on changeable factors such as weather, predation or disease, as on the quality and quantity of the habitat in which the bird must live. If the cover be poor the balance, by necessity, is low; if good. Nature's efforts tend to maintain a population more nearly in accordance with the desires of the sportsman. Furthermore, in either case the crop from year to year may vary widely above and below the average level set by the character of the range. Real abun- dance, like real scarcity, is an abnormal situation to correct which Nature is constantly mar- shalling her forces. Yet, by constant effort, the game manager can improve the habitat so that Nature can strike a higher balance. He can also manipulate many of the limiting fac- tors 90 that over a period of years the balance will be upset more often to the advantage of the sportsman than otherwise. w -^ f. r 5^—re^ CHAPTER XIII FLUCTUATIONS IN GROUSE ABUNDANCE By Robert W. Darrow OCCURRENCE CHARACTERISTICS Amplitude — Rate of Change — Duration of Abundance and Scarcity — Periodi- city — Synchronism CAUSES Relative Significance of Losses During Different Life Periods — Contributory Influences — Predation — Disease — Solar Activity — Weather — Population Density CYCLES SS SUMMARY Grouse fluctuate in abundance in all parts of their range. In addition to the constant oscilla- tion of local populations more widespread fluctuations have been reported at periodic intervals, (p. .556) . With respect to regional areas there has been a tendency for major declines in abundance to recur at intervals of eight to ten years since 1900 at least, (p. 563). Such periods of decline have most frequently occupied two to three years; those of recov- ery three to four years, (p. 560). Periods of general abundance sufficient to afford good hunting have usually lasted three or four years, although longer terms have been experienced, (p. 561). Major declines in grouse abundance during the years covered have not taken place simulta- neously over its range, nor have recurrent declines followed a regular sequence between regions, (p. .567). Pronounced fluctuations at irregular, shorter intervals, often in direct opposition to the gen- eral trend, have been characteristic of local populations, (p. 568). Periodic synchronization of the trends of a preponderance of the component local popida- tions of a region seems responsible for the recurrence of general abundance or scarcity, (p. 571). S56 I Li CTV AT IONS IN GROUSE ABUNDANCE Failure of the annual increment of young birds appears to he the key to such fluctuations. (p. 571). It is not icnovvn what may be the underlying cause of synchronism among such losses. Of the suggested hypotheses, a weather relationship of some kind seems most likely to be con- trolling, (p. 576). Another contributory influence is population density although the nature of its effect is not clear, (p. 576 1. The concept of cyclic behavior is tenable with respect to the fluctuations of grouse abun- dance only if their variability and the uncertainty of prediction are recognized. I p. 579). That wildlife populations are subject to fluctuations in abundance has long been recognized. History is full of accounts of plagues of rodents and insects. With respect to game, as Wing*" has said, in times of plenty "the beknighted savage bowed down in pious supplication and tendered offerings to the Gods and Spirits". During periods of scarcity, however, primitive peoples as well as the pioneers of more civilized races have often had to shift their hunting grounds or be faced with starvation. The, writings of the early explorers and historians record innumerable instances of this kind. Sportsmen, too, although seldom dependent on such species for the necessities of life, have become alarmed when numbers of their favorite game dropped to low levels and have voiced their concern in contemporary sporting journals. But until comparatively recent years such fluctuations were principally of local concern and were regarded as haphazard in occurrence. About the beginning of the present cen- tury, however, observers began to realize that in many instances they had recurred in certain species with considerable regularity. Perhaps the outstanding examples were the Norwegian lemming (Lemmus lemmus) in Europe and the varying hare or snowshoe rabbit (Lepus amer- icanus) in iNorth America. Among the first to call attention to the phenomenon were Mac- Farlane°™, Seton"" and Hewitt"" who used records of the number of hare and other fur-bearer pelts handled by the Hudson's Bav Company to illustrate it. Since then Fllton'^'. '"'. Leopold"' and others have demonstrated fluctuations of more or less regular periodicity in these and a variety of other species over the earth. It has been pdinlrd out in the preceding clmpli'r that grouse populations are in a constant state of oscillation al)i)\e and below an average which is correlated with the quality of the coverts invoKed. Beyond this, apparently more pronounced fluctuations at periodic intervals have been reported. The recurrence, during the past half century, of several quite widespread conditions of scarcity has led lo the suggestion that this behavior might follow a regular pat- tern. It seems appropriate, therefore, to review the evidence brielh and lo discuss how the (lata iif llie present Investigation relate to the problem. OCCURRENCE Many early accounts apparently refer to the occurrence of periods of abundance or scarcity among ruffed grouse populations, but the data are scattered and fragmentary until the latter part of the nineteenth century. It is interesting to note, however, that as early as 1721. the OCCURRENCE 557 species reached such a low level in the vicinity of Quebec that the provincial governor issued a decree prohibiting its being shot"'. Later Nuttall'"', although believing that the birds had migrated, made this interesting observation with reference to November 1831 "in the present season, in travelling nearly to the extremity of New Hampshire, not a single bird of the spe- cies was now to be seen," although he states they had been "so unusually abundant, previously to that period, as to sell in the market of Boston as low as 121/4 cents apiece." Another in- stance is the statement of an old market hunter that a companion, catering to the Saratoga Hotel (Saratoga Springs, J\. Y.), had extreme difficulty in finding birds shortly after the Civil War. More data are available for the years after about 1880, mainly in the form of observations published in various sporting periodicals and in the reports of State Game Commissions. In the more settled sections of the East many such accounts actually referred to the elimination of grouse as a result of man's occupation of the land. Most of these records, however, apply to localities where no serious impairment of the range had taken place. Several compilations of this material have been made. Based on statements in the literature augmented by j)crs()i)al correspondence and inter- views, Leopold"" plotted a curve of relative abundance from lo80-1929 for Vi'isconsin, al- though prior to 1892 it is rather indefinite. The same author also summarized similar data for the north-central states from ]898-1929~'. Michigan records were summed up briefly by Tubbs"" while Clarke'' gathered information for Ontario covering 1873-19.S5 and for other parts of Canada for various periods between 1893 and 193.5. For iSew York and the north- eastern region in general Bump™ presented a preliminary analysis of the trend of grouse abundance covering 1880-1935, though here again the records for the early years are scanty. In considering these compilations one must bear in mind that they represent merely esti- mates of the average grouse abundance over the region involved from year to year on the basis of the material available to the authors. Conflicting reports for different localities in a region in the same year have been frequent. Thus if additional observers had reported the average might have been shifted somewhat. Furthermore the available source material is largely expressed in broad terms so that the comparative degree of change between abun- dance and scarcity cannot be interpreted precisely. As Leopold"' comments regarding his graph. "The vertical ordinate has no numerical validity, the curve simply connecting the most frequent of five grades of abundance reported as obtaining during each year." Nevertheless these records have clearly indicated an alternation of abundance and scarcity, for the period covered, with respect to the general status of the grouse populations of the regions involved. Presumably the same holds true throughout the present range of the species except, perhaps, in environment so marginal that the birds never become numerous. Further- more, since this behavior has been observed among populations situated in remote localities where the environment is still in a primitive condition (Gross'' and others), it may be as- sumed that it has been a characteristic of the species from time immemorial. On the other hand Leopold has suggested that the optimimi portion of the original range of the species lay in the agricultural belt and that fluctuations were less prominent there"'. Unfortunately it is impossible now to test this hypothesis. ' More recently several studies, each extending over a period of vears on the same area, have been conducted. The first of these was the record kept by Criddle"" from 1895 to 1929 at Aweme, Manitoba. Similarly King'"' reported the trend of fall abundance at Cloquet, Minne- sota, from 1931 to 1933. In Michigan, Fisher^'" conducted annual censuses on five areas, the .iS!! ILL CWATIONS IN GROUSE ABUNDANCE longest series extending from 1932 to 1936 although estimates for the Pigeon River tract were continued through 1940 by tlie Michigan Conservation Department.* Finally that of this In- vestigation has covered 1930-1942 and 1931-1942 on its Connecticut Hill and Adirondack study areas respectively, the records of which have been already presented.^ These data also demonstrate the constant fluctuation of grouse abundance. CHARACTERISTICS In order to pursue further an appraisal of these fluctuations the major features of those which have been recorded may be examined. The territory considered has been limited mainly to the range of the grouse east of Manitoba and the Dakotas. and from Pennsylvania north. In this area environmental conditions are most nearly similar to those of New York. Amplitude While it is evident that grouse populations have repeatedly fluctuated from abundance to scarcity and back again it is less clear just what has constituted abundance and scarcity at different times. Only recently have studies in a few localities furnished estimates of density in terms of birds per unit of area. Most of the evidence is of a much less precise nature. As has been pointed out the assumption of similar values for the two extremes in various graphs which have been published covering the past half century has been purely arbitrary and with- out numerical validity. There is no assurance that successive "highs" and "lows" have involved similar densities. Another complication arises from the fact that the season during which grouse hunting has been legal has changed considerably over the yegrs. Throughout the '90s the usual opening date was sometime in September. Undoubtedly such early hunting colored many accounts, of abundance at least, disproportionately in comparison with j)resent experi- ence during the open season. Nevertheless there can be no doubt that the differential has often been great. A few accounts, however, suggest densities which have been involved. Thus in 1881 a writer in Forest and Stream" described grouse as plentiful in Franklin County (N. V.) and stated that one man bought over 2,000 for the market. Another observer reported seeing more than 150 grouse in a day near Lowell, Michigan in 1887*. In 1899 a correspondent at Smithville Flats, N. Y.'"^ commenting on partridge being "more numerous . . . than they have been in a dozen years," noted putting up 50-100 in a day's hunt in September. Again in 1906, just before the general scarcity reported the following year, an item from Chenango County (N. Y.)"" reported flushing 40-50 birds a day and commented that they were "more common than last year." Leopold"' has recorded a number of statements by market hunters and others indicating high levels of abundance in the north-ientral states on manv occasions during the late nineteenth century. Turning to scarcity, three early records have already been mentioned^ Similarly a report from southwestern Ontario in 1883' stated that not a single individual was seen in a whole day's tramp where two years before grouse had been plentiful. Later Carppnter" related that he found only two grouse in the vicinity of Moosehead Lake, Maine from September 27 to October 10, 1887. An even greater dearth of these birds in northern Maine was noted in 1899 by Hardy"" who saw only one partridge in traveling "150 miles by canoe and half as far by foot" during the early fall. Another observer'"' stated that he found from two to eleven grouse near Woodhaven, N. Y. in 1917 where in former years 25 would have been found. In Pine * Ruht. H. D., pcrional letlrr to the aulhori Drrcinbcr 12. 1911. A Sec Cliaptcr XII. p. 539. t Sec diflcuition of Occurrence, p. 556. CHARACTERISTICS 559 County, Minnesota in 1929 King* reported less than two grouse per 100 acres. Recent studies, however, are beginning to furnish a knowledge of what the range of fluctu- ation may be on the same area. Thus King"", although reporting a density of 55.2 per 100 acres on an 1,800-acre tract in Minnesota in the fall of 1933, concluded that an average area of four square miles in the same region would have dropped from about 38 to three birds per 100 acres from 1933 to 1936. In Michigan, Fisher'"' recorded decreases in density on the same scale of from 25.6 to 4.7 (1932-1935 on Pigeon River tract of 2,520 acres), 36.4 to 10.3 (1932-1934 on Munuscong Park tract of 2,044 acres) and 24.1 to 2.8 (1933-1937^ on Houghton Lake tract of 2.574 acres). With respect to the present Investigation it seems evident that the population on the Con- necticut Hill study area in the fall of 1928 was below five per 100 acres. From this point it rose to 21.1 in 1932 and fell to 8.0 in 1912 with several pronounced intermediate fluctua- tions as shown in figure 58. On the Adirondack area the extremes were 9.0 in 1934 and 2.8 in 1935. That densities experienced during recent years have been comparable to what has been considered abundance on many occasions in the past is evidenced by the fact that when the grouse population on Connecticut Hill has reached or exceeded about 15 per 100 acres, it has not been difficult for two men working the more likely coverts to flush more than 100 birds in a day. 100 50 Z < ui 2 O tc u. z o > u o H Z 50 u o I I I 100 FIGURE 57. Connec+icui Hill Ad ir-oodacU. I 100 50 50 '-' 100 1930 1931 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 1942 YEAR RELATIVE DEGREES OF FLUCTUATION OF FALL GROUSE POPULATION DENSITIES ON CONNECTICUT HILL AND ADIRONDACK AREAS — 1930-1942 In this connection it is interesting to note that, while the densities involved on the Adiron- dack area have been much less than those for Connecticut Hill, the relative degree of fluctu- ation has been quite similar in both localities (figure 57). * Id Leopold^^. ^ 1937 populalion estimate supplied by Michigan Conservation Dept. 560 FLiCTUATIimS IN GROUSE ABUNDANCE Hate of Change Of interest also is the rapidity with which grouse aliundaiKC may change to scarcity and vice versa. Considering first areas of regional size, several representative records of declin- ing abundance may be cited. In New York, for example, the last great decline brought the grouse population to a low le\el in 1927 or four years after its high point in 1923, al- though it is important to note that most of the change took |)lace during the years 1926 and 1927"'. Similarly the previous low ebb in this State was reached in 1917'' four years after the abundance of 1913. though the birds were still at a high level in 1914. Here again most of the decrease took place during the last two years of the period. On the other hand the marked scarcity of 1907 followed a high level of abundance in 1906" ". A similar pattern was observed over most of the Northeast, except that longer periods were involved in New York (1914-17 and 1924-27) and Pennsylvania (1914-17 and 1924-28) than elsewhere during those decades. In Ontario Clarke'^ concluded that "such diminutions have occurred ... in the following years: 1933-4 (and 1935 presumed); 1924-5; 1914-6; 1904-5-6". Two or three year periods of decline were reported from the Lake States between 1914 and 1917, the greatest decrease be- ing experienced in 1916 and 1917. In the following decade the years 1924 to 1926 marked downward trends in Michigan and Wisconsin with the sharpest fall occurring in the last two years of the period. In Minnesota, however, an abrupt drop in abundance took jdace in 1924. Then in 1934 and 1935 grouse populations throughout all three states fell abruptly to a low level after reaching high densities in 1932 or 1933*. Similarly the Lake States experienced major declines within one or two years after the high level of 1942. Turning to the rate of recovery grouse reached a peak of abundance in New York in 1935'^, six years after the scarcity of 1927-29, although a high level had been attained by 1934 and in many localities the birds had become numerous by 1932. By 1921 the birds were reported^" to have generally recovered from the low point of 1917 although thev did not reach their peak until 1923, and 1912"" marked the end of the depression begun in 1907. The trend at this time seems to have followed a similar course over most of the New England area. Clarke's interpretation of material from Ontario"' indicates that recovery most often occu- pied four or five years but he apparently included (he last vear of the low period. With re- spect to Wisconsin'"' "' the population had regained a high level by 1921 or 1922 following the low of 1916-17. In Michigan'"' "' grouse had become numerous by 1931, four years after the trend started up in 1928. More recently (19401 |Hipulalii>ns ihronghoul the Lake States generally again became abundant after the scarcity of 1934-36. One may conclude, therefore, that regional trends have usually involved several years and that the period of decrease has averaged shorter tlum tluit of recovery, although both have exhibited considerable variation. The most frequent intervals of decline have been two to three years in length, with tile sharpest droj) taking place during the last year or two. Recov- ery, on the otlii'r hand, has usually been more gradual, three or four years being the most frequent intervals. But a regional population is merely the comjiosite of a host of local populations which may or may not follow similar [)atterns of fluctuation. A knowledge of these local variations is thus of importance also, but unfortunately records of this kind are few. • Prt.oniil lonrm lo the nulhots : Kcrilin.-. C. Dr, rmbit T. 1916: l,r.i|inlil. A.. January II. I1.1S ; Ruhl. H. D.. January 21. I9S5. A I hp ralr iil recovery ta iodicatei) liy ihr niinihrr til grouae rcpnrli'd taken by hunlrts rarli year aince ihr cloaed aeaaona nl 1928 and 1929 ai followa: 1930. 37,188; 1931. 40,448; 1932. 55.845; 1933, 63,910; 1931. 81.614; \t:is. l;«..30l. CHARACTERISTICS 561 Grouse abundance on the study areas used in this Investigation has exhibited both gradual and abrupt rates of change (figure 58). On Connecticut Hill the fall population, after the low point of 1928, climbed steadily upward until a high level of abundance was reached in 1932. Then in 1933 a sharp set-back took place but was followed by an immediate recovery the next year. This was succeeded by a 3-year decline and a 2-year recovery ending in 1939. Since that date the trend has been downward, sharp losses taking place in 1940 and 1942. It must be pointed out, however, that the low points reached in 1933, 1937 and 1940 repre- sented only about half the differential between the levels of 1928 and 1932. Not until 1942, three years after the high density of 1939, did the population again approach real scarcity. On the Adirondack area the population did not exhibit any sustained decline during the Investigation. Rather it reversed its trend almost annually. .Nevertheless it is noteworthy that the year of greatest scarcity (1935) followed immediately after that of greatest abundance* and was succeeded by recovery to a high level the next year. In Michigan, Fisher's data'" show a generally gradual rate of decline in grouse abun- dance on the areas he studied between 1932 and 1936, although in 1933 the population of his Area IV dropped much more sharply than those of the other study tracts while that of his Area III did not begin to decline until 1934. King, however, working in Minnesota in- terpreted his data as indicating a progressive increase from srarcitv to abundance covering seven years followed by a 3-year decline to scarcity again'". Thus it is evident that local populations also have failed to exhibit a uniform rate of change with respect to their fluctuations in abundance. Moreover, although often following a trend roughly in harmony with that of the surrounding region, they have also been sub- ject more frequently to rather abrupt intermediate fluctuations. Duration ok Abundance and Scarcity The foregoing discussion has dealt with the rate of change between high and low levels of abundance. Of interest also is how long periods of abundance and scarcity have lasted. On each occasion, although some one year witnessed the highest or lowest average density reached, there have usually been several years during which the general status of grouse abundance has been either good or poor. The duration of such periods has been from two to four years in most instances. By far the most frequent period of scarcity has been three years. Such periods, of course, include the last year of decline and also the initial year or so of recovery. With respect to general abundance sufficient to afford good hunting, three and four year periods have predominated. Here again an overlap occurs since the first year of abundance corresponds to the last year of recovery. That they may last longer, however, was shown by the 1924-28 scarcity in Minnesota and by the good years of 1939-44 in Vermont and New Hampshire. Furthermore, in case of a minor depression, like that experienced in New York in 1936-37, grouse may be generally numerous throughout such a region for js many as thirteen years or more'''. Periodicity As evidence of the recurrent nature of these fluctuations accumulated, sportsmen and natura- lists alike become interested in their degree of regularity. One of the earliest observers to sug- gest a fairly constant interval was Hatch, whom Forbush"" quotes as saying "that there is an * As mentioned eUe^liere there is some indication tlial the density in tlie fall of 1931 may have been as high or higher than that in 1934 but specific data are lacking. A The current period in New York began about 1932 ami, disregarding the minor depression of 1936-37, had not abated in 1944. S62 I LI CTU AT urns / V GROUSE ABUNDANCE iiiiusuul scarcity of llic birds aliuut once in eight years, and lliat tliis has occurred three times in his cxperieiRe of thirl\ )fars".'" In 1918 Burnham' concluded that, in recent years, shortages of grouse had recurred at about lU-year intervals. Hut it was the scarcity of 1926- 29 that really stimulated study of this phase of the problem, although a sharp loss of produc- tivity in 1924, especially in parts of the .Northeast, had resulted in the formation by the .Amer- ican Game Protective Association of a Grouse Inxestigation Committee and its subsidiary the New England Grouse Investigation Committee. Since that lime a number of workers have com- piled information indicating, for their respective regions, what the trend of grouse abundance has been. The principal accounts of this kind are those of Leopold"', Clarke"*, Bump"", Grange' ', Foote"", Tubbs'"', and Wing"', and, although the data are more or less indefinite prior to the turn of the century, they do reveal that a major decline and recovery took place throughout most of the range of the species during each decade between 1900 and 1940. These regional accounts together with supplemental data, both contemporary and more recent, derived from State and Provincial reports, personal correspondence with State and Provincial officials, wildlife journals, and sporting periodicals furnish a background against which to interpret the data of the present Investigation. In assembling this material, however, it has been found that different statements for the same region often fail to agree regarding specific years. For this reason it has been difficult to determine in some instances just when the high and low points of grouse abundance were reached, although the occurrence of scarcity can be plotted more definitely because it has always caused greater concern. Another feature is the fact that fluctuations have seldom followed a smooth curve. Rather than rising and falling steadily between 1-year peaks and troughs, the trend of abundance has exhibited high and low levels which have usually lasted from two to four years and which have themselves often been characterized by considerable variation. Nevertheless the data can be correlated sufficiently to permit defining within narrow limits the intervals which have occurred in recent years between periods of abundance and of scarcity (figure 59). In New York the records leave no doubt that grouse abundance over most of the State reached a low ebb in 1907"", 1917^ and 1927\ Moreover there is a general agreement in these dates throughout the Northeast, although the decline of the late '20s apparently took place a year or two earlier in New England'^ and the Maritime Provinces', and a year later in Pennsylvania". There is also evidence of less severe reductions in 1903"° and 1924*" and some indication is to be found of scarcity during the late '90s. Omitting these minor "lows", however, there has been a remarkable regularity in the spacing of the other periods at inter- vals of nine to ten years. But since the low period which began in 1927 no comparable widespread scarcity has occurred in New York although there was a minor depression in 1936-37, apparently similar to those of 1903 and 1924. Yet more pronounced declines in grouse abundance were re- ported during the years 1934-36 in other sections of the Northeast'"' '■^. During this period all of New England and eastern Canada were affected. In fact, in 1936 the open season in Massachusetts was curtailed on this account"". \^'here these declines occurred they followed the preceding lows by intervals of eight to ten years. Moreover, the same was true of the minor depression of 1936 in New York. Summarizing his data for Ontario, Clarke'* recognized diminutions among ruffed grouse during the years "19.3.3-4 (and 193.S presumed I ; 1924-.S: 1914-6: 1904-.5-6; 1894-.S; 1883-4-5: 1874" indicating an average period of "between nine and ten years". * SuleniPiil maJe ia 1908. A Pertooal correipoadcnce with State and Proviorial official*. CHARACTERISTICS 563 More recently grouse in Pennsylvania. Maine and much of eastern Canada variously dropped to low levels of abundance in 1943 or 1944* although in Ontario the decline of 1943 was minor. Again the intervals involved were of eight to ten years in length. In New York, Connecticut, Massachusetts, Vermont, and New Hampshire there has still (1944) been no significant general decrease in the present decade. It may be noted, however, that in these states 1944 was only the eighth year since the last depression although, as has already been pointed out, the latter was not marked in New York. In the Lake States (Minnesota. Wisconsin and Michigan), these fluctuations have exhibited an even greater degree of regularity. There, considering only records since 1900. a condition of general scarcity was reached in the years 1907-08, 1916-17, 1924-26, 1934-3.5, and 1943-14 (figure 59). The intei"val involved has varied from eight to ten years with nine the most frequenf^. The fact that so high a degree of periodicity has now been recorded for four suc- cessive lows in this territory seems especially significant. The intervals which have elapsed between recurrent times of high abundance are not so clear as those between lows. The records are frequently indefinite as to just when the grouse populations of different regions reached a high level following the various periods of scarcity. The same is true, although to a lesser degree, with respect to peak numbers. A better criterion, in view of the available data, seems to be the last year of high abundance preceding a major decline. In New York, according to Conservation Department records and other data, 1906, 1914, and 1923 seem to have been such vears. So was 1935. although it was followed by only a minor depression. Thus, in this State, three inter\als of eight, nine and twelve years, respec- tively, have been experienced since 1900. In lite fall of 1944. eight years after 1935. the grouse population over the State as a whole was still at a high level. Elsewhere in the northeastern United States and eastern Canada intervals of eight to ten years have predominated. In much of the East, however. 12-year periods occurred between 1923 and 1935. In Pennsylvania this was followed bv one of seven years associated with the minor low of 1936-37 after which the population fell off sharply in 1943. In New Bruns- wick a short-lived decline occurred in 1940 but the birds were again numerous in 1942 before the scarcity of 1943. Throughout most of this region the records cover four periods since the beginning of the century, but in southern New England, as in New York, only three are involved, although grouse numbers have recently been at a high level for several years. As in the case of scarcity a remarkable degree of regularity has existed in the Lake States throughout four successive periods with respect to the recurrence of grouse abundance. The interval has varied from eight to ten years in length, nine years being the most frequent. These data thus lead to the conclusion that the general abundance of grouse over the major regions of its range in eastern Canada and the northeastern and north-central United States has, during the present century at least, exhibited recurrent fluctuations between high and low levels, the intervals between which have varied from eight to ten years in the case of lows and from seven to twelve years in the case of highs; that in both cases nine years has been the most frequent^: but that, in spite of these narrow limits, there is no evidence of a precise periodicity. * Peraonal corii'si>oudence with State anil Provincial officials. A Leop.ild^sr. suinniari7p(i data taken principally from the north-iciitral states for the year* 1902-27 as indicating two intervals of ten tn eleien years hetween peaks of ahiindance and two intervals, one of twelve years and one of nine years, between troughs. giving; "an average period of a fraction over ten years". But he lumped the several species of grouse together while the data used ahove apply to the ruflcd grouse only. t The means of the records for tlie intervals between high ao.i low abundance as shown in figure S9 are 9.3 and 9.2 years re. speclively. 564 FLUCTUATIONS IN GROUSE ABUNDANCE One must realize, however, the distinction between the above and the inter\'al between the end of one period of general abundance or scarcity and the beginning of the next. The record indicates that in the case of abundance this interval has most frequently been from four to seven years in length while in the case of scarcity it has most often been six or seven years. But this does not mean that there was any need for four to seven year closed seasons. Abun- dance is used here to denote a quite uniformly high population level, while enough birds to justify hunting have usually been present over longer periods with a corresponding shor- tening of the intervals of acute shortage to which closed seasons would apply. So far the discussion of this topic has dealt with the trend of general abundance over areas of regional size. In most years, however, different localities in the same region have exhibited conditions diametrically opposed to each other as well as various intermediate gradations. This raises the question of to what extent the same individual localities have been involved in successive manifestations of this phenomenon. Unfortunately, few specific data based on actual censuses of the same area have been re- corded covering a period of years sufficient for such analysis. Griddle's graph"" indicates a fairly regular periodicity of close to ten years but the area involved comprised less than 50 acres with a correspondingly small number of birds and only the portion from 1914 to 1929 was based on numerical census figures. Furthermore, the latter are spring records and there- fore not entirely comparable with fall data. In Michigan grouse population censuses on the Pigeon River tract of 2.520 acres, begun by Fisher in 1932"' and continued until 1940 by the Conservation Department, showed a steady decline from a high level in 1932 to relative scarcity in 1935 followed by a gradual increase to moderate abundance in 1940. The only other record long enough to be considered is that of this Investigation. In figure 58 have been plotted the fall population estimates for the two principal areas studied. Tlie data for 1930-42 on the Connecticut Hill area and for 1932-42 on the Adiron- dack area (approximately 2.200 acres and 816 acres respectively), represent actual censuses conducted by the Investigation. The approximate trends for several years before and after these dates have also been indicated on the basis of other information. These records reveal that the fall population levels on these areas lia\c fluctuated widelv from year to year rather than following a steady course. That the trough of 1936-37 on the Connecticut Hill tract was minor is evidenced by the fact that the populations of 1927-28-29 are known to have been below that of 1930. Furthermore abundance on this area in 1933 dropped to a level comparable to that of 1937 but immediatelv recovered the next year. Al- though intensive field work became impossible after 1942. the limited data available clearly indicate that the trend continued to decline at least through 1941 so that grouse numbers quite probably fell to the 1930 level if not lower. On the Adirondack area initial work in Jainiar\. 1932. indicated lluit \\m' fall population of 1931 was comparable with, if not high(>r than, any recorded later. The most severe decline occurred in 1935 but was of otdv one \ears duration. Siiue 1936 grouse abundance remained generally high throughout the surve) and sup|)lemental information shows it was still high in 194-1.. In figure 58 smoothed curves* have been superimposed on the actual data in order to indi- cate the general trend which has taken place. These, too, show the trough in 1936-37 on Con- necticut Hill to have been rjiinor and that it was not until 1944 that the population again * Smoothing wan done orcurtting to tlii- (orniula a -4- 2b -4- r. in whirli )i i» the >fiir in (lucHlinn. a llir year brforc and r thr yc»T alter. ^ CHARACTERISTICS 565 Ac+uqI Popula^-ioo EsVrrnotcs . App»-oximate Populotion Levels Before and Af ^-er Pet-tod of Siudy Smoothed Vcalues- 10 O - 300 liJ ■a 200 D Z 100 ADIRONDACK -1 1 1 1. 1928 1929 1930 I93P 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 1942 1943 1944 YEAR FIGURE 58. TREND OF FALL POPULATION LEVELS ON THE CONNECTICIT HILL AND ADIRONDACK AREAS WITH SMOOTHED VALUES SUPERIMPOSED reached a low ebb comparable with that of the late '20s. The interval between these major depressions would thus be at least 17 years. Nevertheless it is significant to note that the minor depression of 1936-37. which came at a time when much of New England was experien- cing a scarcity of grouse, occurred nine years after the low of 1927-28 and eight years before the scarcity of 1944 on this area. On the Adirondack area there was a moderate depression in 1935 but it is not known when the previous scarcity took place. Synchronism The data presented so far show clearly that grouse pojjulations have characteristically ex- hibited an alternation of abundance and scarcity. Furthermore, the records indicate that, at least since 1900, these fluctuations with respect to the average number of grouse over regional areas have recurred with considerable regularitv although their periodicitv has not been pre- cise. In general, during this time, a high level of abundance followed by a decline to scarcity has occurred in each decade. The specific years involved, however, have differed between regions. The record for the north-central and northeastern United States and Canada, as well as it has been possible to interpret it from available information*, is illustrated in figure 59. The range divisions considered have been states and provinces because the principal sources of data have been official reports and other records in terms of such units. In two instances (Massachusetts-Connecticut and Vermont-New Hampshire) states have been grouped because of their similarity and relatively small size. In these cases the records for certain years con- * The data upon wliich this chart is based have been compiled principally from official reports of the several states and provinces as well as from personal letters to the authors from conservation officials in these areas. Use has also been made of the writ- ings of various research workers who have studied the problem. But comments in the literature of purely local conditions have usually been loo few and scattered to be of much value for this purpose. 566 FLUCTUATIONS IN GROUSE ABUNDANCE flict somewhat necessitating interpretation as to the average status of grouse abundance. Ontario, on the other hand, has been divided into three parts according to Clarke' ivhose data have been used for the years through 1934. Admittedly a closer alinement with ecological boundaries would be desirable but the data have not been adequate for such treatment. MINNESOTA O • O • 0« 09 C3» WISCONSIN 0« O* O* O* O* MICHIGAN O* O* O* O* O* NORTHWESTERN ONTARIO O • O • O • O • :;« CENTRAL ONTARIO O • 0« O • O • CO EASTERN a o • O • O • O • •..» SOUTHERN ONTARIO NEW YORK O* O • O • C'O PENNSYLVANIA 0» O • O • O* O • MASSACHUSETTS 8 Q* O • O • 0« CONNECTICUT VERMONT a 0» O • O • 0» NEW HAMPSHIRE MAINE O* 0«0* • • QUEBEC O • • O • O • NEW BRUNSWICK O* ;:« :/» NOVA SCOTIA O • O • 1900 1905 1910 1915 1920 1925 1930 1935 1940 1945 YEAR LAST YEAR OF GENERALLY HIGH ABUNDANCE FIRST YEAR OF GENERALLY LOW ABUNDANCE O Berore major decline • Following major decline '.': B«Fore minor- decline O Following minordecline FICtIRE 59. YEARS OF PRINCIPAL DECLINES IN THE ABIINDANCE OF RIFFEI) tJROl SE IN VVKIOI S RKCIONS OK ITS NOKTIir VSTFRN WP \OliTlI-( FNTU M. R VNCK lOflO-lO]! Two indices have been used in lliir- rliart. First, ii.- lias hoeii luentiuiicil before, the occur- rence of scarcity seems to have aroused the most loiuerii and thus affords the best criterion of the fluctuations which have taken place. For the purpose of this discussion the year i)iotted CHARACTERISTICS 567 in each case has been that during the fall of which the grouse population first reached a sig- nificantly low level even though the absolute bottom of the trough may have been a year or two later. Not only is it often difficult to determine which season was lowest but the year in which scarcity was first reached must have been more closely associated with the conditions which were responsible. The second index is that of the last year of real abundance preceding each major decline in each region. Here, too, the year plotted is not necessarily the one during which the fall grouse population was at its maximum. Rather it marks the end of each period of generally high abundance and, in conjunction with the year when the following low ebb was reached, further delineates the primary interval of decline. It is evident from this chart that the major declines in grouse abundance during the years covered have not taken place simultaneously over its range. Records indicate that the nearest approach to uniformity probably occurred in 1907. although in 1917 also grouse ])opulations over a large proportion of this range were reported to have reached a low level at the same time. But in the latter instance there was wide variation in the year when the decline com- menced. Since then the pattern has been more irregular, exhibiting the greatest degree of variability during the decade between 1920 and 1930, when, paradoxically, the initial year of decline seems to have been quite uniform. Even closely allied regions such as Minnesota and Wisconsin have not usually been affected simultaneously. Yet considerable similarity is shown in the records for Minnesota, Michigan and northwestern and central Ontario. In New York the declines of 1907 and 1917 coincided with similar fluctuations in Pennsylvania and New England, but subsequent years have shown a lack of harmony in this territory. Apparently the populations of various regions tend to react independently of one another. Similar conclusions have been stated regarding the population behavior of other species. MacLulich''" after intensive study of the snowshoe hare in Ontario demonstrated that '"the last year of great abundance, which is the year in which the decrease began" varied from 1932 to 1935 ill different districts. Likewise Cross"' after studying the red fox iVulpes julva) in Ontario found, with respect to different regions of the province, tliat "there were wide variations as to the years in which the maximum numbers occurred, the years in which the minimum numbers occurred and in the intervals between these years". On the other hand, it has been suggested that the onset of grouse scarcity, while not at any time sinniltaneous over the range, might nevertheless follow a similar course from region to region in recurrent instances. The records are conflicting in this respect, however. Clarke's data for Canada in the '30s'' exhibit a trend begiiming in the northeastern and northwestern districts at about the same time and progressively converging southwestward and southeast- ward respectively. A pattern of this kind during that decade is further indicated by the fact that conditions in Maine and the Lake States roughly coincided with those of Quebec and Ontario while the decline in New York. Pennsylvania and southern New England took place still later. Figure 59 also indicates that the pattern was generally similar in the previous decade. But no such progression is apparent in the records prior to 1920. Neither does it seem to be true of the current situation although this has not yet run its course. In this con- nection it may be mentioned that in Nova Scotia the recent period of grouse abundance seems to have ended earlier than elsewhere even though this was not similarly true of the time when the population reached a significantly low level. Thus the available evidence does not support the idea that recurrent declines follow a regular sequence between regions. Yet in 568 FLUCTUATIONS IN GROUSE ABUNDANCE spite of this lack of harmony throughout the range of the species in the onset of major declines in abundance, in most instances there have been a year or two during which grouse have been generally scarce. To analyze the problem further it may be pointed out that the average status of grouse abun- dance throughout a regional unit of range represents the sum of conditions on the many local areas involved. Moreover, while it is obvious that if the average abundance over a region fluctuates the populations of individual areas must also fluctuate, it does not follow that they all exhibit the same pattern. That in actuality they have not done so is evident from the rec- ords in the literature. In these accounts one finds all degrees of variation from agreement to direct opposition within the same region at the same time. As Bump°° commented "it is a rare year in which there does not occur some apparently reliable report indicating both abundance and marked scarcity in the same part of this range". Leopold"'^ in assembling data relative to the onset of the decline of the '20s in Wisconsin, found "a completely irregular incidence, such as moths would make in invading a carpet". Simi- larly Green and Shillinger"" observed that pojiulations "may become abundant at the same time and die off at the same time in widely separated areas, whereas in two closely adjacent areas an abundant population may develop in the one while animals* are dying off in the other". In correlating the available information for Ontario and other provinces in Canada. Clarke'' found "local differences of at least three years in the time at which diminution begins". The same author, in describing his method of summarizing questionnaire information'' indicates that reports of both abundance and scarcity were received from the same region during most years. Gross'" recorded a parallel situation for New England. In 1937 Connecticut" reported "islands of abundance in a fairly general scarcity". A great many other accounts also attest the fact that such dis])arity among local grouse populations lias been of frequent occurrence. During the most recent jjcriod of severe grouse shortage in New York (1927-28) Allen" reported that the birds were "still apparently in normal numbers" in a few places in the cen- tral and northeastern parts of the State. In fact the ]jo])ulatioii of one compartment of the Connecticut Hill study area is known to have been high in 192f). Likewise W oodruff"' quoted a State game protector to the effect that in Allegany County (N. Y.) in 1907. when grouse in the Northeast generally were at a very low level, "Partridge seem to be more numerous than last year — fifty birds seen a day". During the present Investigation the grouse populations of the two princi])al study areas have followed similar courses during some years and at other times have been divergent (fig- ure ,S8)^. In 1931. while the Coinieclicut Hill area was still recovering from the scarcitv of a few years before, the numl>cr of birds on the Adirondack area was at as high or higher a level as at any time after that. From 1932 to 1939 both areas exhibited generally similar trends althr)ugh the low point reached on the northern tract in 193.'i was relatively much lower than the depression of 1936-37 on Coimecticut Hill. Since 1939 tlie trend on the latter area has been steadily downward while this has not been the case on the other. In this connection it may be noted that the minor trough on (^.onnecticut Hill in 1936-37 coincided with the status of grouse reported at that time for New York as a wlnilc iiiul the Northeast in general. Fisher's data for Michigan"' afford another example of a lack of synchronism because, al- * The tenn animal wat used here as applyini; to both the {rouse and varying hare. A "The etrpss o! the percentage rrportlng 'abundant* over those rcporlinf 'scarce* was the figure used in plotting the graphs.** T It is reroRni/ri) that llicse imto arfiis difTrr holh in sire atiit in rhararler, the Ctmnri'lii-ul Hill trmt hring madr up <>( a num- ber nf separated coverts while on the Adirondack unit. the gniuse cover is continuous. In the absence of more extensile data, however, their comparison seems warranted. CHARACTERISTICS 569 though they corroborate a general shift throughout the north-central region from high to low numbers between 1932 and 1935 followed by gradual recovery, they show that individual areas were often at variance with one another. For example, in 1933 when populations on the Pigeon River and Munuscong Park tracts were found to have decreased sharply over the preceding fall, that of the Houghton Lake area more than doubled. Two years later abundance on Munu- scong Park took a marked upturn while it continued to decline at Pigeon River although pre- viously the two areas had paralleled one another. These and many other records serve to demonstrate that ultimate understanding of the part synchronism plays in the fluctuation of grouse abundance must rest on a knowledge of what happens on the smaller units of range. The data of this Investigation for its Connecticut Hill study area offer a clue to the nature of this relationship even though no condition of severe scarcity was experienced. D«nsi+u v/ilhir) comparlmenT Tr«nd of densily -for whole araa 50 Z ^ 100 o It ll. z g < > U 0 o I- z u o 5 50 0. 100 I I I I I 150 100 50 '/ \ / /• / / : i ■ • I \* ■ ' . : ' 1 I I 50 LJ 100 1930 1931 1932 1933 1934 1935 1936 1937 1938 1939 1940 1941 1942 YEAR FIGURE 60. RELATIONSHIP OF TREND IN THE AVERAGE DENSITY OF THE FALL GROUSE POPULA- TION ON THE CONNECTICUT HILL AREA TO THE DENSITIES WITHIN THE VARIOUS COMPARTMENTS COMPRISING THE AREA As has been discussed elsewhere* this area is not a continuous unit of grouse habitat but comprises a group of woodlot coverts more or less separated from each other by open land. The basic portion, which has been used for appraising the population trend of the area as a whole, has been divided into twelve compartments of 130 to 300 acres each largely * See Methods and Techniques, p. 695. 570 FLUCTUATIONS IN GROUSE ABUNDANCE conforming to these individual coverts. The grouse populations of these compartments be- haved quite indejiendently of one another and seasonal estimates of numbers were made for each. The sum of these constituted the estimate for the whole area. In figure 60 the fall population density on each of these compartments has been plotted year by year in terms of its percentage deviation from the mean for that compartment together with the trend of that for the area as a whole. It is significant that there was a wide spread in the degree of deviation among the different units each year and that each fall some units were at a low level although the specific ones varied constantly. It is also significant, with respect to those compartments which were low in various years, that the degree of deviation from the mean was often as great when the aggregate population level was high as when it was low. In fact in 1942, when the population as a whole fell to the lowest point since the first year of the study*, all but two compartments were within the same range of deviation that had been observed during other years. The diflereme lay in the fact that in 1942 a greater pro- portion of all the units were "below the line". Similarly years of abundance were those in which the coverts producing an above-average number of birds outweighed the ones below average, but nevertheless each year witnessed some units below average. It is important, too, to realize that the relative rank of the different compartments, as shown in figure 60, constantly varied from year to year. To give some idea of the lack of unison observed a composite graph of the individual trends has been prepared (figure 61). 400 300 O QC O 200 tc hi d D Z 100 I I I I 400 300 200 too 0 '-' FIGURE 61. i_ -i- -L. _1_ 1930 1931 1932 1933 1934 1935 1936 1937 YEAR 1938 1939 1940 1941 1942 TRENDS OF FALL i;ROLSE POPULATION DENSITIES FOR THE VARIOUS COMPART- MENTS OF THE CONNECTICUT HILL AREA — 1930-1942 Thl■^il■ data, although covering only a comparali\cl) siiorl pci ioil of time with respect to tlie behavior of wildlife populations, seem to demonstrate that the abundance of grouse in iiidi- \i(hiai coverts is conslanllv fluctuating and that, even during seasons when the birds are plen- tiful over a tract as a whole, many such coverts are normally below average. It seems prob- * FraKnirntary data iniliratr thai t)ir populalinn on lliin arra was aomewhal lower atill in 1943 and 1944 but 1942 it the Uat year fur which (lata comparable to the other year* of the Itudy are available. CAUSES 571 able that a similar relationship exists with respect to the status of grouse abundance through- out any range area. The data also suggest that conditions of general scarcity which occur from time to time over wider areas may be primarily a consequence of some combination of factors causing a loss of productivity over a preponderance of the component coverts, but that the actual populations of many of these coverts may often fall little lower than they do in other years. Synchronism, with respect to the status of grouse abundance over any range area, thus seems to have been greatest during recurrent major troughs, but to have occurred a year or so later than many parts of the area initially reached a low level. Considerable lack of syn- chronism has been characteristic of the early stages of most declines. The term "spotty" has been applied to conditions of this kind when grouse are still plentiful in many localities but noticeably scarce in others. Lack of synchronism has also been frequent during times of generally high abundance. CAUSES The reproductive potential of a species constantly tends to increase its numbers. The potential productivity of the ruffed grouse is greatly in excess of that which can actually be realized because the range will accommodate only about so many. For this reason low popu- lations tend to rise, but those already at a peak have little o|)|)<)rlunity to increase and tend to fall back. A wide variety of influences operate as limiting factors. Most of these vary from year to year, and even from season to season, causing grouse populations to fluctuate continually in numbers. Moreover, radical changes in the environment may cause local scarcity at any time. The observations of this Investigation as to the manner in which these influences have affected grouse productivity on the areas studied have been discussed in the preceding chap- ter. These data have shown that differences in the degree of survival during one or more life periods rather than changes in the reproductive potential of the birds are the chief reasons for the population changes recorded. It is probable that this is typical of the behavior of such populations. It has also been shown that, in addition to the constant but irregular fluctuation of local populations, there have been recurrent periods of general abundance and of general scarcity over greater areas. It has been shown, too, that these periods, particularly those of scarcity, have recurred at intervals of eight to ten years in many parts of the species' north-central and northeastern range during the time since 1900. Furthermore, data have been presented which suggest that such longer-term fluctuations in the average abundance of grouse over extensive areas may be primarily a result of the recurrent synchronization of a preponde- rance of the local populations at low average levels, although at such times many of the com- ponent units exhibit densities little, if any, lower than are often reached in other years. In other words it seems evident that the immediate causes of these major fluctuations in general abundance lie among the same agencies which limit survival on local areas regard- less of to what extent they may be governed by more deep-seated influences. Relative Significance of Losses During Different Life Periods The data increasingly point to the success or failure of the annual increment of young as the key to the problem. As already discussed under Net Productivity*, this has been the most frequent cause of low fall populations on the Investigation's study areas although just when * See Chapter XH, p. 539. 572 FLUCTUATIONS / V GROUSE ABUNDANCE the loss occurred has varied on different occasions. On Connecticut Hill losses during the breeding and nesting season have averaged proportionately greater than those during the brood period. In 1940 high nest mortality caused a sharp drop in the fall population level. On the other hand, in those years when failure of the crop was most severe (1933, 1935 and 1942), unusually high brood mortality always occurred. On the Adirondack area very high brood losses were responsible for the poor crops in 1933 and 1935. Although breeding fail- ure apparently occurred its significance is not clear. Unfortunately there was no opportu- nity to observe conditions associated with a real scarcity throughout the State. Evidence that failure of the ainnial increment has frequently been the chief cause of major declines in the general abundance of grouse is also found in the observations of others. Among the first to report this were Woodruff*"' and Forbush'" with reference to the scarcity of 1907. As a result of his investigations in New York. Woodruff noted that "with only one exception the observers state that all or nearly all the birds shot in 1907 were old birds, and most of them cocks. Scarcely any young birds of the year were seen." The same author also quotes similar reports from Connecticut. Massachusetts and Michigan. Speaking of New England, Forbush stated "I had been much in the field during the spring and summer of 1907, and had noted that very few young birds were reared in the region with which I was familiar." Also applying to the 1907 decline the following statement appeared in the Annual Report of the Pennsylvania Game Coramission™" "Game birds have not been as plen- tiful during the last season as heretofore, notwithstanding the promised increase because of the many birds left over the year before." Regarding the decline of the following decade Minnesota stated"" "In 1915 all hunters re- ported finding few coveys of young birds and the few found were small. Much the same condition prevailed in 1916." Similarly Pennsylvania's Annual Report for 1916"™ commented that most of the grouse killed in the fall were old birds. Again in the next decade an official report for Minnesota"*' contains the following state- ment, "During . . . 1923. the increase in partridge, or ruffed grouse, in many districts of the state was almost unbelievable, these birds abounding in great numbers in suitable lo- calities. So far as it has been possible to ascertain the birds wititered well during the winter of 1923 and 1921 but for some unaccountable reason at the close of the nesting season in 1924 the decrease in numbers of these birds was so enormous that the closing of the tKirnial shooting season was being seriously considered." In northern Michigan, too. a large crop was said to have hatched in 1924 but most of the young died before the hunting season"'. Similarly Brewster" called attention to "the well known fact that every now and then almost no young grouse arrive at maturity in Now England during seasons when very many are successfully hatched". McDonald'", speaking of eastern Canada, reported a great abundance in the spring of 1924 and a good hatch but few at the end of the summer and also com- mented "In fifty years with the grouse I fiiii)rte kind lia\e been carried on by Green and Shillinger* and Clarke"' as well as by this Investigation. Grouse have been found to be subject to a great variety of diseases and parasites. Outstand- ing among them as decimating agents of wild grouse are the stomach worm ( Dispharynx), tularemia and a blood ])arasite ( Leucocytozooii ). The last of these in |)articular was found to be associated primarily with mortality among chicks. Yet none of these maladies has been found widely distributed throughout grouse range, while those parasites which have a wide (lislriliulion appear largely to be innocuous'^. On the other hand, the fact that there is still so little known of the paramount causes of brood mortalily leaves the possibility that some important but as yet unrecognized disease exists. But a.'-suniing that sonic coiulilion of ibis kind ucic loiind. a m((lKuii>Mi capable of svn- ilnonizing its irruption o\er large areas and al the same time allowing for e\len>i\e local \ai'ialioiis would remain to be discoM-red. Clarke' has suggested what might lake place if Leucocytozooii were llw principal cause involved. i?ut the presence of Leucocytozoon has * See varioui paper* by Green and ShiUinter. A See Chapter X, p. 4-12. CAUSES 575 not been demonstrated in connection with grouse fluctuations outside Ontario. Nevertheless, whatever may be the underlying cause of widespread diminutions in grouse abundance it seems probable that it is usually accompanied by lowered vitality among the young. If this is true, the stage would be set for disease to add momentum to the decline even though the specific organism involved might be different in various regions. Solar Activity As students continued to ponder the geographic extent of recurrent major fluctuations in grouse abundance as well as their seemingly high degree of harmony they began to look for some all-pervading force as the cause. About this time interest was stimulated in the pos- sibility that variations in solar activity might be correlated with such behavior in wildlife pop- ulations. Using sunspots as an index a relationship between solar activity and terrestrial life, presumably through the medium of rainfall, had been suggested in data on rates of tree growth"". DeLury"' and later Wing"" reported fluctuations in bird migration dates which appeared to correspond more or less to changes in the number of sunspots. Elton'" inter- preted records representing the trends in abundance of several species of birds and mammals, notably varying hare and lynx, as following patterns sufficiently close to the 11.2 year peri- odicity of sunspots to indicate a connection. Here seemed to be the answer — a force capable of causing the continent-wide sweep believed to characterize grouse declines. But it has become evident that such a relalionshi]). if it exists at all. is not as simple as at first envisioned. As brought out at the Malamek Conference"" most iSorth American spe- cies, for which the available data indicate some regularity in the intervals between recur- rent fluctuations, have exhibited periodicities of less than ten years which do not correlate with that of sunspots*. MacLulich" later demonstrated this quite conclusively for the varying hare and lynx (Lynx canadensis). Similarly the records for ruiled grouse, to the extent that they represent periodicity, have exhibited an a\erage interval of 9+ years which cannot be directly associated with the sunspot interval of 11.2 years". Furthermore the data of this Investigation and other recent studies, as well as more critical analysis of older records, have shown far less uniformity in the fluctuations of the component populations of a region than was once thought to be the case. Nevertheless no one can deny that the sun is the original source of energy upon which the existence of life on the earth depends. Changes in the amount of effective solar energy reaching the earth, although relatively small, are known to take place. Moreover, sunspots are but one manifestation of these changes. Some force not yet recognized may be involved. But, however that may be, there seems little doubt that, if any connection does exist, the motivating mechanism with respect to synchronizing fluctuations in grouse abundance must operate quite indirectly and be subject to considerable variation. Several intermediate factors which might function in this way have been suggested. One possibility is that the virulence of disease organisms might vary with changes in solar ac- tivity; another that the vitamin content of food or some other nutritional factor might be affected; still another that some influence associated with climatic or weather conditions might be responsible. Yet little evidence has so far been presented for the first two and even the degree of relationship between weather trends and changes in solar activity is still being debated"' '"• ^"'- "'^. * Leopold'^^ has shown a lack of agreement also for the red grouse in Great Britain. A Grange'^, using Wisconsin data, has suggested a method of weighting sunspot figures in order to correlate them with grouse abundance but it does not provide for differences such as have occurred between various regions. 576 FLUCTUATIONS IIS GROUSE ABUNDANCE Weather Regardless, however, of what governs it, or to what extent it may or may not be related to the forces represented by sunspot changes, the evidence is becoming increasingly strong that weather is a primary factor in governing variations in grouse abundance. Data from the Con- necticut Hill study area indicate that both temperature and precipitation are important. Other elements of weather are undoubtedly involved also. Despite its evident significance, however, a great deal still needs to be explained regarding the specific ways in which weather functions in affecting grouse productivity. With respect to roajor fluctuations, the data discussed in Chapter VI* with respect to the relationship between March and June temperature trends in New York and recurrent grouse scarcity in this State reveal some agreement in point of time. Furthermore, tempera- ture records for other grouse states exhibit similar degrees of agreement. Perhajjs this was coincidence for exceptions also occurred. Certainly the channels through which it may have exerted its affect are not clear. Then, too, temperature is only one element in the complex of factors, even with respect to weather, operating in any instance. All of them vary in intensity and no one of them is likely to be the best index at all times. Nevertheless, the possibility of a connection warrants serious consideration. To speculate regarding the modus operandi of such a relationship is hazardous. One pos- sibility, though, is indicated by observations during this Investigation. In view of the fact that the evidence points most strongly to failure of the increment of young as the key to grouse scarcity of this kind, weather may function through some physiological efifect in which the weather conditions involved would not cause direct decimation but rather would set the stage for subsequent losses. It must be emphasized that this is mainly conjecture. A sugges- tion of something of this kind, however, is found in the data from the Connecticut Hill study area^ which indicate a correlation between temperature and precipitation during the three weeks immediately following hatching and the brood mortality occurring during the latter part of the sununer. In this connection, too. may be cited the observation that a lack of sharp temperature changes from day to night ma) iiihiliit the mating of sheep and thus result in failure of the lamb crop^". If weather should prove to be a significant factor in recurrently synchronizing grouse scarcity throughout large areas, the fact that, in spite of general conditions, it varies greatly from place to place because of such influences as irregular topography and shifting storm tracks would help to explain the disparities between localities which have been so frequently re- corded. ]}ut how much fire there may be under all this smoke must await further research. I'opiilntion Density Aiiiilher piece which must event iialK be fitted into the ])uzzle pertains to the role of pop- iilalinn deiisilv in relation to Hue luatioiis in grouse abuiulafHC. As was discussed in the chap- ter dealing with produc ti\ il\ a IciiiliricN was iiolcd during llic Iinestigation for high breeding populations to be less pruductixc. relatively, than low ones and fur this to result from a lower increment of young being produced". Moreover, on llu- Ciniiici licut Hill study area those compartments which exhibited relatively high densities at limes when the general average was low were often those whiili. for one reason or another, had been low the preceding year. • Sec figure 23. ji. 305. A See Choplei VI. p. 303. t See Clupler XU, p. S40. CAUSES 577 ^^Tiile no severe scarcity was experienced during the Investigation, this, nevertheless, suggests that the immediate causes of such shortages may be effective primarily with respect to high breeding population densities. It also suggests that they will result in general scarcity mainly when they occur in conjunction with a preponderantly high level of abundance among the component local populations of an area. If this should prove to be the case it would help to explain why abnormal weather, such as that of 1912 for example*, was not associated with a pronounced low in grouse abundance"^. In other words it may be that density is a necessary catalyst. Other observers have reported similar experiences. After stating that, although apparently scarce early in the season, grouse in most localities turned up later, a Connecticut report™ continued "However, in places where birds had been very numerous in the previous year, they appeared to be very scarce in 1924." A news release of the Wisconsin Conservation Department in October 1933 stated regarding mortality in certain northern counties in the fall of that year "Up to the present time it has affected ruffed grouse principally, and on areas where the birds were most plentiful." Commenting on a general slump of more than 50 per cent throt|ghout most counties of both peninsulas of Michigan in the fall of 1934 as compared with 1933 Ruhl* stated "In other counties .... characterized by scattered coverts which are heavily hunted, grouse were almost unanimously reported to be more abundant than in the autumn of 1933." Similarh. in speaking of the decline in Minnesota in 1942 and 1943 Blair' re]5orted that it "a|)parentlv was not as effective in the areas where birds were most hunted as it was in the places off tlie beaten trails where less hunting was done." It may be, therefore, that reduction of high population densities through hunting for sport, regulated in accordance with actual ])roductivitv from year to year, can serve to lessen the severity of recurrent scarcity. In this connection it is possible that the lack of niaikcil fluctualioM in numbers reported by Leopold""' for the remaining grouse habitats of the Middle West during recent years, and which he suggested might indicate that the species did not originallv exhibit such behavior in that part of its range, is not an inherent characteristic but rather a result of the fact that the birds no longer attain high densities in that region. It is evident from the foregoing that, while the record concerning major fluctuations in grouse abundance suggests a considerable degree of regularity in their recurrence, there is still a great deal to be explained regarding the mechanism involved. If anything is certain it is that the interaction of several factors is usually responsible. Nevertheless, the data pre- sented seem to narrow the problem to some extent. They indicate that the key lies among the forces which cause failure of the annual increment of young, but that a preponderantly high level of abundance among the component unit populations involved may be a neces- sary prerequisite to the synchronization of the effect of these forces. They also indicate that the rate and degree of decline are closely related to the relative size of the increment com- pared with the breeding population and to the average life span of the species. Further they suggest some of the factors which may implement these relationships and point to weather as of primary importance among them. The remainder of the solution must await further research. * See figure 23. p. 305. A Assuming ihat the temperature faotor eiled also proves signiBrant. t Ruhl, H. D,, jiersonal letter la the authors, January 2J. 1935, t Blair. F. D„ personal letter to the authors, February 28, 1945. 578 FLICTUATIONS /A GROUSE ABi\DA.\CE CYCLES No discussion of fluctuations in wildlife abundance would be complete without including the concept of cycles. Ever since observers began to realize that the interval? between suc- cessive periods of abundance and scarcity of many species had exhibited a tendcncv toward regularit) , the idea of their recurring in cycles has appealed to scientist and sportsman alike. It has appealed particularly to those interested in game because it suggests the possibility that the future course of abundance can be predicted. For this reason it is important to "look at the record," especially with reference to grouse, to see how strongly existing data support this hypothesis. The concept of cyclic behavior is not new. Since the begiimings of time the cycle of night and day, and the annual cycle of the seasons have been recognized. During the past two centuries, however, cycles have been ascribed to a great many other things. Probably the best established, aside from the movements of various celestial bodies, is that of the changes in the sun represented by sunspots. Closely associated with this are the variations in terrestrial magnetism, while less well correlated are fluctuations in ultra-violet radiation from the sun and climatic changes'. But cycles have also been described in such things as the price of common stock, building activity, tlie prciduction of pig iron and death rates from various diseases'". With respect to wildlife, abundance data have been interpreted as indi- cating cyclic behavior among a variety of species chiefly in the northern hemisphere. Among them, in addition to the ruffed grouse, may be mentioned mice and lemmings'"' '"', varying hare and lynx^'", the arctic fox (Alopex lagopusJ''\ the red grouse (Lagopus scoticus) in the British Isles"" and the Atlantic salmon (Salino salar)'". A significant feature about these fluctuations, however, is the variety of periodicities rep- resented. For example, the ruffed grouse seems to have exhibited an average interval between peaks and troughs of slightly more than nine years*. MacLulich"'" concluded that those of the varying hare and lynx were both 9.7 years and Elton'" found approximately the same peri- odicity in the records of the Hudson's Bay Company for the muskrat (Fiber zibelhicus), red fox, marten I Maries americana) , wolf (Canis) and mink (Muslela vison). An interval of be- tween nine and ten years has been reported also for the Atlantic salmon""". On the other hand, mice, lemmings (mainly Leininus trimucronaliis) and the arctic fox iiave followed 1- year trends in North America and Europe'^'- '". Tluil of liie red grouse in Britain has av- eraged 0..5 years"", while its cousin, the willow grouse or rype I Lagopux lagopus) of Scan- dinavia, has shown a pattern of three to four years'". The sunspot cycle has a mean of 11.2 years, while agricultural productivity in the I'nited States, which would sccni to reflect climatic changes, has shown a 18.6-year fluctuation'"'. But. irrespective of the periodicity involved, none of these trends has exhibited a precise recurrence of its successive peaks and troughs. The degree of variability in this respect has differed considerably between species, in man) cases being quite wide. The intervals seem to have been most nearly constant among the mice and Irnmiings as well as the arctic fox. Other species have varied to a greater extent. i"oi ihc lulled grouse from seven to twelve years have elapsed between "highs" in different instances and from eight to eleven years between "lows". This frequency has ranged from eight to eleven years for the varying hare and from eight to twelve years for the lynx. In the case of the red grouse in niilain it has Naried from four to eight years. But even the reconiic! Ilmlualions in >uris]>ots have shown a variable recurrence of from seven to seventeen years. • Sec (liscuaaiiin <■( (•.■ri.iili. ilv, p. .Id:!. CYCLES 579 All this raises the question of just what the term "cycle" should be considered to mean. In discussing the characteristics of cycles in connection with weather phenomena Marvin°'° wrote: Mere recurrences of natural phenomena without reference to the time interval between recurrences do not necessarily constitute cycles. To be cyclic there must be systematic recurrence ... If, as some claim, the variations observed in length and amplitude of the cycle — such, for example, as the differing lengths and intensities of the sunspot cycle — are not of an accidental character but are controlled by law. then the proponent of such a view nmst not only prove the reality of the climatic cvcle he claims, but the nature and reality of the systematic fluctuations in length and intensity must be fully set forth . . . Proof of cycles by verification of predictions is the ultimate criterion, but its utilization is obviously beset with difficulties. If climatic features are the composite of the operation of several cyclical effects then past features must he capable of syn- thetic reproduction from cyclical elements, thus affording proof of the cycles." The same analysis could be applied equally well to the use of the term with respect to the fluctuations in abundance of grouse and other wildlife. If a definition such as the above were adhered to, grouse and many other species could not be classed as cyclic on the basis of existing data. On the other hand, the fluctuations of grouse, for example, have shown a distinct tendency toward rhythmic behavior, especially in certain regions. Furthermore, if accurate quantita- tive data were available, the degree of variability might be found to have been even less. In view of the absence of such data the possibility remains that the trend has followed a sys- tematic pattern and. to prove or disprove it. highly accurate records will have to be kept for some time to come. The data marshalled by Elton'"' with respect to mice, lemmings and foxes afford some idea of what is needed for such a purpose. Likewise, correlation or lack of correlation with the fluctuations of other species or events will require long series of data in each case. This was shown by MacLulich"" who found that the relatively accurate records for the varying hare in Canada* for the 87 years from 1847 to 1934 were insufficient to demonstrate their lack of correlation with the sunspot trend although this could be proven through the use of similar records for the lynx extend- ing over 174 years (1751-1925). But it must be enqihasized that mere correlation of the periodicities of two trends, even though quite perfect, does not mean they are related with respect to cause and effect. For instance, mice near Ithaca, New York were found to exhibit a cycle in harmony with that of Presidential elections'"". Some link connecting the two must be identified. For the present, however, the element of variability cannot be plotted in advance. The next "high" or "low" may occur a year or two before or after that representing the average interval. Amplitude also may vary, as happened in New York in 1936-37, so that a seg- ment of the cycle is largely obliterated. Furthermore, local populations have often been out of step with the general average and the relative status of such populations is subject to change at any time in response to changes in the local environment. With these limitations in mind, continued application of the cyclic concept to grouse may be warranted. But predictions regarding the prospects for fall abundance in any particular year cannot be made with any degree of exactitude on the basis of data assembled to date. Rather such prospects, for the time being at least, must be judged during the same season on the basis of a knowledge of the spring population level and of the degree of breeding and rearing success. * Principally tliose of the Hudson's Bay Company. Nucleus of Grouse Habitat PART III Managing thk Grouse Crop GROUSE MANAGEMENT By Gardiner Bump Game management is the process of putting together known facts to produce, maintain and use the game crop. The days when this had to be done largely "by guess and by gosh" are gradually becoming a memory. In their place, for species after species, research is beginning to supply a blue- print; management to translate, and implement this to produce the desired results. The Investigation, therefore, has sought to marshal the pertinent facts about the ruffed grouse which could be used as a sound basis for grouse management throughout the State. In New York, on the Conservation Department rests the responsibility for fixing, each year, the hunting regulations for grouse. In addition it is called upon to manage some eighty thousand acres of game lands, most of which represent grouse cover. It also attempts to make some provision for game management on approximately half a million acres of submarginal farm land bought for reforestation purposes and on which grouse and other species of game represent a valuable, secondary crop. At the same time there are millions of acres of grouse cover in private ownership. More- over, an ever-increasing proportion of this is finding its way into the hands of people who place a high value on recreational op|)()rtuiiities which such lands can provide. These people seek, and are entitled to have, advice in the development of their land to improve and main- tain a game as well as a forest crop. New York is now, and probably always will be, pri- marily, a forest and woodlot game state. Among the small game species, none is so highly prized as the rutled grouse. Conse(|uenll). its encouragement, by applying appropriate measures on State or private lands, is highly desirable. Management, to use the vernacular, is the "pay-off" on the investment in research. Inevi- tably, this pay-off will come in installments over the years. There are a number of reasons for this. First, it would be presumptuous for the authors to assume that thev have found out everything worth knowing about grouse and have completelv mined the accumulated data for every conceivable productive application. Secondly, while many of the findings have been translated into practical suggestions which have been field-tested, sufficient time has not elapsed to permit full evaluation of all the results. Application, too. is dependent ujjon interest in the problem which bids fair to grow with the years. In the preceding chapters on research the authors have made every attempt to meet scien- tific standards to the best of their ability. In the ensuing chapters, in attempting to translate the findings and the personal experience gained into management suggestions, this will not always be possible. Here, in order to make the most complete contribution, it is often neces- sary to state the case as we now see il. In doing this it is understood that, in the light of future experience, modifications are inevitable. 582 GROUSE MANAGEMENT To encourage a broad understanding of the problem and still present in workable sequence the specific steps by which a grouse crop may be encouraged, the management material has been organized as follows: 1. A brief account of the experimental grouse management work already undertaken on State game lands. 2. A resume of the ten factors affecting the abundance of grouse by way of showing which may be utilized and how in producing more grouse. .3. A discussion of what mav be done on a broad scale to encourage grouse over the State as a whole (Chapter XIV). 4. A detailed, step by step, analysis of how to design productive grouse coverts and set up long-range management plans therefor (Qiapter XV). 5. A discussion of how this may best be carried out by improving and maintaining grouse cover (Chapter XVI). 6. A consideration of other ways of maintaining the grouse crop (Chapter XVII). 7. A discussion as to how best to coordinate the work of various Federal, State and local agencies with private enterprise in carrying out grouse management projects (Chapter XVIII). Experimental Management Projects The bases for the management suggestions have been the present factual study of grouse life history and requirements and the experience and observations of the authors in applying the principles here learned to State game management lands over a ten-year period*. As a part of the testing program, the composition and type arrangement of the cover on each management area were surveyed in the field, then placed on large scale maps. The vis- ual presentation facilitated an analysis of cover conditions forming the basis for subsequent improvement projects, as well as for long range management plans. These were drawn up in considerable detail. The improvement work fell largely into two categories, establishment of cover and its main- tenance. Under the first, open lands were planted to conifers and hardwood species, so arranged as to produce ultimately a substantial timber crop meanwhile providing excellent food and shelter for grouse. Around many of these, shrub borders, utilizing such species as thornajiples, dogwoods, viburnums, apples and evergreens, were set out in clumps (with due 'regard to future hunting possibilities.) In the entire program about 2,000,000 evergreens and tOO.OOO broad-leaved trees and shrubs were set out. Overgrown land improvement projects centered mainly about the problem of maintaining the status quo. This was done by employing cutting, poisoning, and infrequently, burning to arrest the normal succession and eliminate undesirable species. Occasionally conifers were planted in such areas to provide needed shelter. In wooded areas, the composition and arrangement of cover were altered by judicious thinnings and by selective lumbering of merchantable trees. Where forest cover was exten- sive and unbroken, small areas were clear-cut, in rotation, to provide necessary summer and fall feeding grounds and thereby increase materially the carrying capacity for grouse of such cover. In all, about 160 of these were established. <^ Ilii" *».i» iDittrrinlU liiillx-iiil li\ ijir i(«r nf nv.-i ,1 niilliun null (l;iv« ■•( Civilinn Con»ervaliiin f.iirj"* ami rfl>f labor. J GROUSE MANAGEMENT 583 But trees and shrubs, like constructive ideas, take time to grow. Some results of these cultural practices may, therefore, not be fully evident for some years. To bridge this gap. the Investigation also noted the effect, on cover, of cutting, planting, burning and grazing carried on at some previous date in many coverts and under a variety of site conditions. Fitting the Factors Controlling Abundance Into the Management Picture Before considering the management suggestions that follow, it might be helpful to analyze objectively the factual blueprint research has produced. Its overall design embodies the ten factors that, collectively, determine grouse abundance*. Three of these, the physiology, the reproductive capacity, and the general habits representing the basic characteristics of the species are but little subject to man's control. Yet all are adversely affected by insufficient food and shelter which we, within limits, can influence. Another, weather, while in itself un- controllable, can also have its adverse effects mitigated by assisting Nature to provide the proper environmental setup. Disease, so far as can now be determined, is of particular importance only when grouse have been allowed to overpopulate their habitat. A part of the treatment lies in adjusting the yearly harvest so as to leave in the coverts, breeders suflRcient to produce a satisfactory crop for the following year — and no more. This only can be done if the responsible public agency (in the case of New York State, the Conservation Department) has the power to es- tablish the proper seasons, the technically trained men to determine what they should be. the protectors to enforce them and the backing of an interested and an informed public. In addition to this, food and shelter again are important, for a favorable composition and ar- rangement of cover produce healthier birds and tend to distribute them throughout so as to make disease less likely to reach epidemic proportions. Predation is more complicated. When grouse are few, every bird taken by predators, as well as by any other cause, may well represent a significant loss. If the reverse is true, a con- siderable number of birds may fall prey to predators without seriously affecting the general level of abundance. Normal yearly losses from predators, as recorded by the Investigation, have averaged about 40 per cent of the adults. Yet the elimination of such destructive species would not result in saving, for the sportsman, most of these birds unless they were killed in their first fall. Of all those that prey on grouse, onlv the goshawk makes grouse a significant part of its diet. The others catch a bird or break up a nest occasionally, but their main reliance for food is on plants or on other animals. The abundance of these "buffer species." then also influences the part predators play in limiting the numbers of grouse. It is neither possible nor desirable to eliminate all grouse predators from the coverts. On the other hand, in New York State, if the numbers of great-horned owls, foxes and weasels, where they are overly abundant, could economically be controlled, the harvestable surplus of birds would probably be increased. But such control is not practical over large areas. Since losses from predation adversely influence the total grouse population much less in productive than in poor coverts, habitat improvement assumes increased importance. At the same time it seems probable that, on small areas to be managed for intensive shooting, the control of the principal nest predators would result in a greater fall crop of grouse. This would be worthwhile, however, only if the additional birds were harvested for sport. * These are physiology, reproductive capacity, general habits, food, shelter, weather, predation, disease, man, artificial propagj tion. 584 GROUSE MANAGEMENT Of all the factors, man, today, probably leaves the strongest impress on the grouse. But his interests are so diverse, his activities so complex and the results so varied as to make generalizations exceedingly difficult. As a sportsman, his annual kill, when a shootable surplus exists, takes for sport a sub- stantial proportion of the crop which otherwise would be lost to predation or would disperse. Furthermore, where the birds are very abundant, hunting mav function advantageouslv by reducing the population density before disease has had a chance to l)econie important. Con- versely, in a period of grouse scarcity, hunting may slow down recovery by subtracting from the population many birds that might better be left to breed. There is. though, some evidence to indicate that the annual kill, during periods of substantial decline or increase, affects the trend of abundance much less than is generally believed. As a trapper (and hunter as well) man tends to reduce the abundance of [iredators, par- ticularly if they are present in considerable numbers. Much more important, however, are the farmer and the lumberman, for they create, con- trol the development of or destroy the habitat itself. Plow, cow and axe determine the size of the grouse crop in New York State over a period of years to a far greater extent than do trap and gun. Today's best grouse coverts largely result from the abandonment of poor farms and from small woodcutting operations. To subtract these influences from the picture would be to sacrifice, after a period of years, the variety of vegetation and the interspersion of cover without which grouse can never become really plentiful. It follows, then, that those who wish more grouse must, in the long run, encourage such activities or be prepared to pay the cost of the cultural operations necessary to maintain similar conditions. Here is where man's role as a conservationist enters in. No longer is it sufficient merely to regulate hunting pressure. One must also learn to produce and maintain environments in which grouse will thrive. To accomplish this requires organized effort on the part of a public informed on the problem and on the benefits to be derived from solving it. A closely related factor is the ])racticability of n'.slorkinii dpi)leted coverts with hand- raised birds. These are. at best, costly to raise and difficult to produce in any numbers. Nor is there, to date, any indication that, with properly constituted coverts, it is necessary unless the seed stock of birds has been completely eliminated. Throughout this section the emphasis on providing the proper food and shelter is inescap- able. Accordingly in the management section, emphasis has been placed on how to design and manage grouse coverts and on how to maintain tlic crop of liirds. The liroad picture is out- lined in the first chapter uilli intensive management being discussed in tlie succeeding pages. The final chapter deals witli conrdinating and utilizing the forces that large!) must be relied upon lo carry out the work. Coordination nf all f>f the above mentioned forces should result, over a period of years, in maintaining a population Ie\el that will provide a large shootable surplus during most years. It mav ameliorate but will not |)re\enl. however, vearh niiclualions in gmuse abun- dance in any inilixiiinal covert. Our goal. then, sliouki be to build U|> and inainlaln as nian\ high-producing coverts as possible so thai in atn one \ear tbei<' vvill alwavs be an abnndani-c of these carrying a shootable surplus. Tluis reads the formula for nuin\ a niemoiable dav afield. CHAPTER XIV MANAGING GROUSE AREAS By Gardiner Bimp WHY MANAGEMENT IS NECESSARY INTENSIVE VERSUS INCIDENTAL COVER MANAGEMENT THE GOAL — IDEAL COVERTS For Grouse — For Nestings For Broods — For Adults For the Grouse Hunter OVERALL COVER CONDITIONS, TRENDS AND NEEDS IN NEW YORK STATE The Adirondack Region — The Catskills — The Rest of the State SOME PRACTICES HARMFUL TO GROUSE THE ROLE OF THE STATE Annual Responsibilities — Yearly Inventory — Regulation of Hunting — Long-term Projects — Research — Service to the Landowner — Acquisition and Development of Grouse Lands — Maintaining Hunting Grounds — Stimulating Grouse Production on Private Lands — Stimulating Interest and Understanding — Emphasis on Fundamental Conservation Proarams & SUMMARY With constantly increasing hunting pressure on grouse, either the hunter must harvest less or he must see that more and better coverts are produced. I p. 587). Intensive cover management for grouse alone can seldom be justified on a large scale (p. 587). -The management of woodlands for grouse may be combined with the production of other forest products so successfully as to make the real question one of individual interest rather than of opportunity or economics. (p. 587). To better define the goal — the establishment of as many productive habitats as possible — ideal coverts for nesting, for broods and for adults are described, (p. 589, 593). It is perfectly possible to produce coverts providing excellent conditions both for the grouse and for the hunter. Edges, small slashings and openings, properly arranged, are the major prerequisites, (p. 594). 586 MANAGING GROUSE AREAS Overall cover conditions, trends and needs in a heavily-forested region (the Adirondacks), a forest and farm area (the Catskills) and in a farm and woodlot section (Rest of the State) are described, (p. 596). Loiijr-lime maintenance of good cover is, in large measure, dependent on moderate though widespread lumbering, grazing of overgrown lands and stable farming communities. The wide adoption of better forest and soil conservation practices is of definite concern to the sportsman as a practical means to this end. Other suggestions as to how sportsmen and other conservationists may assist are presented, (p. 598). Practices detrimental to grouse cover include clean farming, heavy grazing of woodlands, un- controlled burning, and too much or too little lumbering, (p. 599). Leadership in planning and in carrying out broad projects of benefit to grouse must come from the State, (p. 601). On the State rests the responsibility for deciding on yearly hunting regulations in accordance with the picture provided by an annual census of grouse abundance and conditions, (p. 602). To assure future grouse crops, the demand for which is constantly increasing, the State must also vigorously prosecute certain long tenn projects. Included among these are man- agement research, application of the facts thus found, maintenance of hunting oppor- tunities, encouragement of landowners to produce better coverts and the stimulation of an active interest in grouse problems, (p. 602). #, Broadly speaking, those interested in improving conditions for grouse fall into one of two groups. There are the individuals and organizations who are active in supporting any pro- gram for the betterment of the bird, but seldom have the opportunity for carrying on any actual management work. There is a second group, equally interested but who also own or control grouse coverts and who wish specific guidance in making conditions therein as nearly perfect as possible. This chapter is written with the first group in mind. The others will find it informative as a background for the more specific pattern of cover improvement contained in succeeding chapters. Perhaps the general picture may best be presented by anticipating broad questions, the answers to which will help lo better define the path to be trod in bettering conditions for this grand game bird. Why is management necessary? How intensive should it be? What are we aiming at in terms of an ideal covert for grouse and for the hunter? In general, what activities are likely to result in improved coverts and. therefore, should be encouraged? \\ lial practices are likely to be harmful? What part ran the State play in improving condi- tions for grouse? l.el us consider these, briefly, to avoid confusion, but in detail suiricient to stimulate thoughtful consideration. WHY MANAGEMENT IS NECESSARY •V It is axiomatic that the grouse is a product of the en\ ironment in which it lives. Li'ft to he own devices, Nature only here and there produces a combination of conditions sullicieiitly INTENSIVE VERSUS INCIDENTAL COVER MANAGEMENT 587 favorable to result in a bumper crop of grouse. Yet the number of sportsmen wishing to harvest this crop is constantly increasing. The alternatives are clear. Either the hunter har- vests less or he must give Nature a helping hand by encouraging her to produce more and better coverts. Though of less importance, the local control of the more destructive preda- tors*, and a complete harvest of the surplus of grouse each year, will also help to maintain the supply. But, fundamentally, grouse populations are a reflection of covert quality and abundance. The surest way to have more grouse, then, is to improve the coverts and increase their number. This requires management. INTENSIVE VERSUS INCIDENTAL COVER MANAGEMENT Considering the State as a whole, grouse are but one of several desirable crops, to the production of which wooded areas are adaptable. Each crop has its own SQt of requirements which parallel or conflict at points with those of the others. To fully meet these for any one usually entails an expense greater than the value of that particular crop. On the broad front, therefore, it follows that any management practices favoring grouse must be exten- sive in character, and dovetailed wherever possible to meet the needs of as many other crops as possible. Furthermore they must be modified to some extent where serious conflicts arise. When one considers the individual covert, however, the decision must be made at the out- set as to whether grouse is to be the primary crop or merely an important subsidiary product to be skillfully encouraged as opportunity arises. In many cases, the latter will be the case for few have as yet developed grouse shooting preserves where the sport is so superior as to encourage the expenditure of substantial funds for grouse improvement alone. In fact, the management of woodlands for grouse often may be combined with the production of other forest products so successfully as to malie the real question one of individual interest rather than of opportunity or economics. For instance, in New York State, grouse require a mixture of hardwood and coniferous cover. To encourage such a combination is likewise good forest practice. Grouse need brushy edges, overgrown lands or small slashings to provide summer and fall feeding grounds. Scat- tered through most woodlands are areas, which because of poor site conditions or the char- acter of the present stand, are of little value for the production of timber. With a little thought and effort many of these can be made or maintained so as to meet grouse needs. Here i| should be clearly understood that such areas must be scattered through the wood- land in such a way as to make a large number of good habitats meeting all the predominant requirements of grouse, rather than concentrated in a few locations. In general, if one is willing to devote from 10 to 20 per cent of the total wooded areas to this end, fair to good grouse crops may be had, with very little sacrifice of forest products. In the Northeast where, at present, only a small fraction of our forest lands are producing at anywhere near capacity, there is, therefore, little economic justification for not including grouse production among the principal ends to be encouraged on managed woodlands. It is well to point out that in developing a woodland for grouse many subsidiary ends, each desirable in itself, are attained. Both bank and sheet erosion are controlled, stream flows are regulated, floods made less severe, fire made easier to control. Soil fertility is im- proved to a greater degree when the forest cover is varied than when it is composed of but * No practical method of predator control, applicable on a state wiile basis, is known. 588 MA\A(;i\G GROUSE AREAS one or two species. In the same \\a\ insi-ct ilaniajic lo tree* normally is more severe where the stand is relatively ])ure. Recreational o|)|)orlunities are increased, not alone because more wildlife is present, but also in accordance with the |)riiiciple that variety is the spice of life. Yet there are an increasing number of organizations and individuals who wish to make the production of a grouse crop a primary end in itself. W hile. as a matter of economics, this is seldom feasible, in terms of total enjoyment per dollar expended there is much to recommend it. Perhaps a happy medium might be attained were one to devote 25 to 30 per cent of the covert to fully meeting grouse needs and the rest to producing other forest crops on a self-sustaining basis. This in general is the background against which succeed- ing chapters dealing with the intensive development of grouse areas have been written. "> "^' \ .^N EXCELLENT COVEKT IiN W HK.ll .\ll.\i;i) WOODS ARE EDGED WITH OVERGROWN FIELD.s .\M) BROKEN. HERE AND THERE. BY SMALL SLASHINGS AND ABANDONED LANDS. SUCH A COVElfT MAY PROVIDE LARGE CROPS OF BOTH GROUSE AND TIMBER. IF SKILLFl I.I.V MAINTAINED THK (;OAL IDEAL COVERTS Before considering, even in a general way. what ma\ be done to improve grouse coverts. naturally one nuist have a clear ])icture of the prodiiil desired. \\ hat. then, are the char- acteristics of an ideal covert? Grouse cover must serve the needs of the birds during the nesting, the brood and the adult periods. Since the production of a liarvestable crop is the goal of most game managers, at- tention must also be given a covert's ''huntability." to coin a term. Hnl first, what con- ditions are ideal for the bird? THE GOAL — IDEAL COVERTS 589 For Grouse For Nesting Typical nesting cover exhibits a more definite pattern than one might imagine. The sites chosen are usually favorable, not only for that purpose but also for the youngsters which, when hatched, will need to travel but a short distance to find food and shelter. For New York State the picture shapes up as follows: — Basic Patterns. Second-growth woodlands intersected by woods' roads and bordered by overgrown fields, old pastures, young slashings or patches of popple, birch or alder provide ideal nesting sites. Should a part of the woods be made up of mature trees, it would make but little difference for a considerable proportion of the broods spend the first few days, after hatching, in such a location. If, here and there throughout the second-growth woodlands, a few trees have been cut. perhaps to furnish the winter's wood supplv and thus encourage small islands of briers and other open land vegetation, the area is doubly attractive. Some birds will choose such sites and thereby be somewhat safer from predators than as though their nests were located in the more open hardwoods. Composition. Hardwoods are ])referred as nesting cover although the particular species seem not to matter. While few nests are found in coniferous stands, it is inuiialerial whether or not an occasional evergreen is present in the vicinity of the nests. Undergrowth Density. Little attention needs be paid to providing protected nest sites for the majority of the birds choose a rather open spot against the base of a tree or stump for this purpose. Slope. The nesting cover might cqualiv \\r]\ be located on a steep slope, a gentle iiulinc or on flat lands for nests are equally apt to be located on anv of these. Extenl. The nesting cover need not be extensive, for nests are commonlv placed within 100 feet of an edge. Woods' roads or open trails within the forest seem equally attractive in this respect. For Broods Optimum brood cover in New York State is characterized by its youth and its diversitv. It is the early stages of woodland succession, in which are found fresh herbaceous growth in considerable profusion, that are attracti\c. Here, then, are the most common brood cover patterns and characteristics: — Basic Patterns 1. Second-growth woodlands surrounded by broad bands of woods' edge type vegeta- tion preferably fairly open in character. 2. Second-growth woodlands interspersed with substantial amounts of overgrown lands, old shrub-dotted pastures, brushy corners or brier patches. 3. Moist alder runs, separated by hardwood slopes inclined to be dry in summer. 4. Small slashings, still in the brier, herb and small sprout stage, scattered through second-growth woods. 5. Selectively-lumbered hardwood stands in which the openings thus created are filled 590 MANAGING GROUSE AREAS z o s z o 3 •J f- o ^ b. til r. la] K o ^ 5^ u Q o < ^ ^ 05 o t/: t^ z a o c/ ^ z o i^ O o ^ O s- S Id < a U U u O o 2 HJ O s i o H < z u U O 3 z e: J O 3 m o ^ — : ^? ^ ./¥ ± .*:■-' H-'T^P^- *; -s *.,. ' ^rf J -rWv --»^«wjfc._f'' >5 ""^V /. .** ;j^ m ^:0- ..'i'^,. "*s^m^M:M^ THE GOAL — IDEAL COVERTS 593 with vegetation characteristic of recently cut-over areas, though usually somewhat less varied and abundant. Grassy spots are seldom much used. Composition. The greater the variety of the vegetation, the better. Conifers are not impor- tant and predominantly coniferous woods are but seldom utilized. Density. Woodlands rather open or of medium density are considerably used, especially about the edges. Overgrown lands and slashings, which are preferred, may be densely vege- tated if rather open spots are scattered throughout. Undergrowth. Variety here is likewise important. Small hardwoods, shrubs, berry bushes and lush herbs are preferable. Slope. Steep slopes are avoided. Water. Open water, such as a stream, is not a necessary component of brood cover since dew, succulent vegetation and moisture-fdled fruits meet brood needs in this respect. Indi- rectly, moist spots, because they encourage a profusion of herbaceous vegetation and insect life, are quite attractive to broods. For Adults The hallmarks of quality in cover for adult grouse are variety in composition and a high degree of interspersion of the cover types. Woodlands, particularly if they contain some conifers, are the backbone of a covert although brood cover, brushv patches and cut-over areas are much used, particularly in summer and throughciut the earlv fall. To meet year- round needs the ideal covert should include the following: — Basic patterns. Typical cover consists of patches of second-growth or mature conifers scattered through a stand of mixed hardwoods and conifers, edged by second-growth hard- woods* abutting overgmwii land or ciil-over areas. For the sake of clarity, let us further subdivide this into the following seasonal patterns: — Spring — Second-growth hardwoods adjacent to coniferous clumps or to mixed hardwoods and conifers. Summer — Slashings, spol-lumbcrcd woodlands and inalurc or second-grow ih liardwoods. Conifers are not desired. Fall — Broad bands of rather open woods' edges, overgrown lands or slashings adjacent to a mixed woods of hardwoods and conifers. Winter ■ — Coniferous patches scattered through second-growth hardwoods or mature hard- woods and conifers. Com position. Pine, hemlock, sjiruce. l)alsam and white cedar all provide excellent shelter. In mature stands, an undcrslory of mixed coniferous and hardwood reproduction is desirable. A substantial part of the hardwoods present should be species producing food for grouse. Birches, poplars and cherries are excellent in this respect. Beechnut mast, when available, is eagerly sought after. Small beech gro\ es or wide-spreading old trees scattered throughout the woodland make for more attractive cover. Density. Excej)! as heretofore indicated, the density of the woodland cover seems not to be directly of prime importance. Clumps of conifers are, perhaps, most used when of med- * Important primarily for neetins cover and sliould not bt- extensive. 594 MANAGING GROUSE AREAS ium density although the relatively very dense cover, typical of many reforested areas, may be much frequented for a few hundred feet in from the edges. Hardwood stands of medium density are most satisfactory. Undergrowth. The important point here is to have a variety of undergrowth available beneath the crown cover. The density is of less moment though extensive patches of thick undergrowth are not particularly productive while contributing materially to the difficulties of the hunt. Large areas covered with grass or rank herbs such as goldenrod or asters are usu- ally avoided. Slope. Neither slope aspect or degree are particularly important insofar as adult grouse are concerned. Water. As with broods, a stream or other open water is not a necessary component of ideal adult grouse cover. The above represents a broad pattern towards which one may direct his efforts. But. though the basis lies in an analysis of 19.619 adult grouse flushes. 1.515 brood contacts and 1,270 nest locations, it is unquestionably possible to deviate from it to some extent and still produce many grouse. For the Grouse Hunter A much more controversial subject is the planning of a covert to provide not alone favor- able conditions for the birds but for those who would hunt them as well. Here the authors have but little to go on* save their own experience as enthusiastic and persistent grouse hunters, and those of other experienced sportsmen who have generously contributed their ideas on the subject. Naturally the first specification for an ideal hunting cover is that there be at least a fair concentration of grouse. This varies in most coverts with both the year and the season. One may expect but will not find a good crop every year even in the best of coverts. To be reasonably assured of good hunting opportunities, one must, therefore, have a number of excellent coverts from which to select those with currently high populations for hunting. Even where birds are usually abundant, every hunter knows of some coverts which are seldom hunted because the birds do not "fly right" when flushed. What he usually means by this is that food and shelter are so intermingled that, though many birds may be flushed, they seldom remain in sight long enough to provide even a sporting chance for a shot. It is perfectly possible to have highly productive coverts which also offer excellent hunting oppor- tunities. With respect to the hunter, in the Northeast, at least, an ideal covert is one which con- tains a number of edges in which the birds are apt to be found feeding, especially during the early part of the hunting season before the leaves have fallen. No grouse enthusiast need be told of the attractiveness in October of hedgerows, old pastures and overgrown lands, espe- cially if they adjoin a woodland of mixed hardwoods and conifers. Woods' roads lush with clover, also, then, are much frequented. Later, the birds may be expected to concentrate more in the wooded areas, particularly if evergreens are present. * A diicustioD of the lyppa of covrr to Khii-h icroiiftr atr tiinsi likely In fly vt\\vn fliislicd. will \tv found in Chaplrr III. [>. 166. ^'- * THE GOAL — WEAL COVERTS 595 Another favorite feeding spot throughout the fall and early winter is the cut-over land or slashing. Small slashings scattered throughout the interior of a covert, especially if bor- dered by conifers on at least one side, provide a likely place in which to find birds when they are not feeding in the overgrown lands. It is desirable also to have the covert bisected or at least deeply indented with one or more narrow openings across which some birds are likely to fly when flushed from the adja- cent cover. Favorite hunting coverts usually provide these conditions in abundance. But there are also other considerations. Thick cover seldom adds to the enjoyment of the hunt, nor do steep slopes. For the former reason, slightly grazed woods and overgrown lands often furnish the best hunting possibilities. Dense, extensive blocks of conifers or large tracts of hard- woods without evergreens are equally undesirable. Here the cover is either too thick to shoot or too poor to attract many birds. For hunting, an ideal "edge" in New York State may be made up of clumps of trees and shrubs, including thornapples, apjiles, viburnums, dogwoods, cherry and evergreens growing in a relatively open setting though immediately adjacent to a rather open mixed woodland. Here a dog works well, and the birds, once flushed and heading for the woods, usually must cross one or two small openings, thus providing superlatively good shooting opportunities. The size of the cover types that make up a covert is also imjiortant. in that extensive areas of overgrown land or of mixed hardwoods and conifers tend to scatter the birds more and to make them diflTicult to follow once flushed. These jxiiiils are illustrated in figure 62 where the cover has been so organized as to pro- vide not only excellent conditiims for grouse but for the hunter as well. SCALE I •660 LEGEND : OVERGROWN FIELDS .'■K"-i SECOND-OROWTHHjgn HARDWOODS a!]^ HARDWOODS B^liU AND CONIFERS lllll:! CONIFERS 77Z\ SMALL cZZA SLASHI FIGURE 62. ONE OF MANY POSSIBLE ARRANGEMENTS OF COVER TYPES WITHIN A COVERT WHICH SHOULD PROVIDE EXCEPTIONAL CONDITIONS FOR BOTH BIRD AND HUNTER 596 MANAGING GROUSE AREAS OVERALL COVER CONDITIONS, TRENDS AND NEEDS IN NEW YORK STATE Considering the State as a whole, grouse cover is made uj) of a vast assortment of indi- vidual coverts. The extent to which these, individually and collectivelv. fall in with the pat- terns heretofore descrihed. largely determines our grouse crop over the years. The improve- ment of individual coverts specifically for grouse is important. But there are also broad influences and activities at work which, though often not recognized as seriouslv affecting the qualit) of the grouse range as a whole, yet exert a substantial influence upon it. Like- wise, there are certain characteristics of the land and the use man makes of it which must be taken into consideration. To better picture these, let us outline the situation and needs in a heavily forested region (the Adirondacks) , a forest and farm area (the Catskills) and a farm and woodlot section (the rest of the State). The Adirondack Region* Grouse cover in the Adirondack region is characterized by extensive, often unbroken forest areas. Only the larger valleys are farmed. Most of the region lies within the Adirondack Forest Preserve of which 36 per cent (2.1 71. .538 acres) is in State ownership. On this no improvement activities for either forest or game are permissible, for under the constitution of the State it must be kept as "wild forest land forever." This provision effectively pre- vents exploitation of the timber resources. It prohibits not alone lumbering, but also any betterment of the existing cover for timber or wildlife production. As the forest grows and matures, it tends to become more even-aged and less broken up by openings, thus making conditions for grouse relatively unfavorable. Interspersed with State-owned lands are private holdings. These are generallv extensive. Many are in the hands of absentee owners. Periodically, most such lands are cut over in large blocks for lumber or pulp. As a result, except about the periphery or in the vicinity of farmlands, conditions are generally not particularly favorable for grouse and large concentrations are seldoin to be found. Such conditions, quite frankly. |)lacc marked limitations on wiiat nia) be done to improve the grouse crop. Any aclivit) whiih tends to diversify and break up the extensixe forest areas into smaller blocks is. for the birds, a move in the right direction. The application of sound forest improvement and lumbering practices to private lands, and the location of more small wood-using industries within the Adirondacks would niateriallv advance this objective. Those interested in grouse in this region would also do well to encourage, wherever pos- sible, farming and pasturing since overgrown lands and attractive woods' edges are largely byproducts of these activities. More intciisi\e development, except on a small scale, is scarcely practical unless grouse i)ecome an important subsidiary crop worthy of attention in its own right as a part of the recreational opportunities for which this section of the State is famous. The Catskill Region In the Catskill region, witli il> tairiis and \alleys. pasliiicd >lopc> and wooilcd uplands, the situation is (piili' difFerenl. Though nnich of the area lies within the Catskill Forest Preserve. State ownership is much less extensive. Moderate pasturing, coupled with a large number * A gtrnrrjll ili'si-riplion i>( i-nih ri-pion t» In be found brginninc on p. 111. OVERALL COVER CONDITIONS, TRENDS AND NEEDS IN NEW YORK STATE 597 of usually small lumbering operations, has tended to diversify the cover and maintain pro- ductive edges to a much greater extent than in the Adirondacks. These, coupled with the progressive abandonment of the poorer or less well situated farms, has resulted in the estab- lishment of much fine grouse cover. Conifers are generally less abundant than in the Adiron- dacks, however, and good winter shelter, particularly in the more extensively forested parts, is often lacking. In this region, as in the Adirondacks, the maintenance of a good grouse crop over the years is intimately tied up with the adoption of good forest and soil conservation practices and the continued encouragement of small lumbering and farming operations. \^Tiere grazing is heavy, cattle should be fenced out of the woodlands. Fencerows should be encouraged and stream banks, gulleys and steep slopes allowed to revegetate naturally without interference by grazing. The planting of small blocks of conifers in old fields and pastures is another activity worthy of considerable encouragement. The Rest of State Region Except for the Lake Plains, the Lower Hudson Valley and Long Island, the rest of the State is characterized by well-farmed valleys lying between poor uplands. Here one finds the largest areas of good grouse coverts in the State, for nuii'h of the land, once cleared, has been worked to exhaustion and abandoned. Overgrown fields, old pastures, long forgotten or perhaps still moderately pastured, seeding in to apple, hawthorne, pine and cherry, are the rule. Many woodlots, too often and intensively lumbered over for their own good, provide an abundance of summer and fall feeding grounds, though the preference for softwoods has resulted in the elimination of much desirable winter shelter. Between lowland and upland, such a wide diversity of conditions exist that this region may well be subdivided into fanned lands and largely abandoned uplands. On the former, condi- tions are not unlike those described for the Catskill region. In general the farmer, with cattle and axe. is maintaining a fair amount of ovei^rown land, hedgerows and pastures. Wood- lots are lumbered upon occasion to pro\ idc for the normal needs of the farm, .'^niall timber sales are common. Here the grouse hunter has a definite, tliough often unrealized, stake in any program which directly or indirectly helps to maintain farming activities on such lands. Over the years, the price of milk and wool unquestionably will exert a stronger influence on grouse than will the limit of the hunting season, for cow and sheep are still the most effective instruments for slowing down the natural reversion of j)astures to forest. In like manner, to encourage the wide adoption of better farming and forest standards and practices, calculated to improve soil and forest conditions, is to help provide a practical, though indirect means of main- taining present coverts for the future. On the abandoned uplands, the situation is quite different. Cover diversification, so impor- tant to grouse, is here a fact. Open fields are being rapidly swallowed up by woods as brush lots become second growth forests in a surprisingly short period of years. Large tracts are being reforested with conifers, either privatelv or bv the State. The problem, here, is how l>est to maintain the diversity of forest and overgrown cover since but little of the land is pastured and lumbering is sporadic for necessity, born of poverty, thoroughly denuded the woodlands of merchantable timber before abandonment. To sharpen up the picture, let's be more specific as to what will have to be done under each 598 MANAGING GROUSE AREAS of the conditions described above, to maintain grouse cover. On the farm lands, to help hold back the forest and maintain about it lliat fringe of over- grown fields so important to grouse, one might well encourage the following activities: 1. Those programs which help the farmer to maintain his economic positions. Included in these are those of the Soil Conservation Districts, the State Forest Practice Act. the farm extension program of the State College of Agriculture and the interest in rural electri- fication. 2. Dairying and the moderate pasturing of sheep as an indirect means of maintaining the overgrown pasture. TVPICAI, ABANDONED UPLAND On the abandoned uplands, to maintain an indispensable diversity of woodland cover, a more active interest must be developed in projects such as: — 1. The wide acceptance of forest standards as set U]) under the State Forest Practice Act. 2. The selective cutting of woodlots. 3. More local wood-using industries. 4. The fencing out of catllc from woodlots. 5. So managing woodlands as to stimulate coniferous reproduction in predominaMtly hard- wood stands. 6. The encouragement of a wide band of woods' edges about woodlands. 7. A program of reforestation of idle lands ])roviding for: — (a) The |il;iiilln;: of eonifers in small blocks. (b) Variety in the s|)cei<-s to be planted. (c) The eventual develo|imi-nl of stands composed of hardwoods as well as soft- woods. (d) The lca\ing of wiilc borders between blocks and of frequent openings within SOME PRACTICES HARMFUL TO GROUSE 599 them which are allowed to grow up to trees and shrubs producing food for wildlife. While the importance of maintaining good grouse habitats and increasing their numbers cannot be overemphasized, the picture would not be complete without reiterating the desir- ability of continuing other types of activities. Chief among these are: — 1. The wise regulation of hunting so that surplus populations may be harvested or hunt- ing pressure reduced in accordance with the situation. 2. The hunting and trapping of the more destructive predators, particularly those which break up grouse nests. 3. The stimulation of an intelligent, vigorous interest in conservation in general and in grouse in particular. To accomplish some of these objectives will require a re-orientation in thinking and activ- ities, even among those who have the best interests of the grouse at heart. It is human nature to live for the present. This being here, wise heads will look to the future and direct today's programs into chaimels which will ])rodu(e more grouse tomorrow. SOMK PRACTICES M AHMFUL TO GROUSE Many an activity of benefit to grouse, when carried on in moderation, may have the o])po- site effect if pushed to extremes. Wildlife, because of its more or less specialized require- ments for survival, has suffered particularly from the exploitation, beyond reasonable bounds, of such ideas as the draining of swarn])s and clean farming*. It is. therefore, worth consider- ing for a moment some practices that may ad\ersely affect the grouse. All tend to standardize rather than to diversify covert \egetation. Among them should b<' niciitioncd the follow ing: — 1. Clean farming 2. Heavy grazing of woodlands 3. Uncontrolled burning of woodland and overgrown fields 4. Large lumbering operations within individual forest tracts 5. The prohibition of lumbering 6. The complete reforestation with conifers of large contiguous areas 7. The maintenance of relati\ely pure stands of any species of tree, shrul) or herbaceous growth The undesirable aftereffects of the first three are too widely recognized to need further emphasis. With respect to lumbering, too little or too much, concentrated in any one area, is apt to encourage uniformity in cover. A slashing half a mile across will be used largely along its edges. The removal of all the merchantable conifers from a woodlot. seldom leaves conditions as attractive as they were even though berries follow the axe. Acid-wood cuttings are particularly damaging in this respect because of the frequency with which whole hillsides are reduced to briers and sprouts. Where, however, as in parts of the western Catskills, the trees are removed in long, narrow strips, each separated by second-growth woodland, the situation is reversed and more favorable conditions are the result. * The elimination oi hedgerows and the pasturing of steep slopes anil gullies are examples of this. 600 MANAGING GROUSE AREAS Conversely, the policy of sparing the axe and letting Nature take her course is certain in the long run to produce forests replete with cover furnishing winter shelter but sadly deficient in summer and fall feeding grounds not alone for grouse, but for many other species of wild- life as well. • In principle the situation may be much the same where conifers are planted in large blocks, unbroken by brushy openings, hedgerows or hardwoods. Only along the edges will such plots be much used by grouse. On the other hand, properly thinned and opened up to encourage a mixed stand of hardwoods and conifers rather than a dense, lightless evergreen mass, a plantation can be quite attractive to grouse. By leaving strips, along hedgerows, old apple orchards, thornapple clumps, swales and similar cover furnishing food for grouse, unplanted much can be done to provide vegetative variety. Most plantations lack this from the time the tree crowns meet until opened up again bv thinning or lumbering operations. AN IDEAL ED(;F. SUCH AS THIS IS OF MORE VALl'E FOR H U.DI.Il K THAN AS A SITE ON WHU M TO PI.VNT EVERGREENS In considering reforeslalidn one must remember lluil it is an acti\it\ aimed piimarilv al restoring idle, run down acres to a productive .status. Kor this |)urposc there arc sound rea- sons for planting conifers in close rows. So long as this practice is followed, there will be periods in which the trees are growing up when grouse will be found largely along the edges, uidess openings arc ))rovidcd within. Once the crop trees become merchantable and cutting is begun, however, conditions for grouse are bound to improxe. F.ventualh, uidess the forest is clear-cut and replanted again to conifers, the composition ma\ be expected to approach that of the surrounding woodlands. The suggestions, given in llic preceding paraiiiaph sim- ply tend to hasten this day. THE ROLE OF THE STATE 601 With regard to the last mentioned harmful practice, fortunately, one seldom finds conditions in the Northeast so uniform as to stimulate large areas composed almost exclusively of one or two tree species. Where these do occur, either naturally or as a result of some activity controlled by man, grouse usually are notable by their absence. Common sense and scientific fact are combining to counsel moderation with all of these practices. They are their own worst enemies, in that when vigorously carried out they tend to so far destroy the balance of Nature as, ultimately, to build up a strong resistance to the very things they seek to accomplish. To combat them it is necessary first clearly to establish cause and effect by pertinent research, and secondly, to discourage the practice by emphasizing the undesirable results. Only by such a program, even though it cover a span of years, can many of these harmful practices be combatted. THE ROLE OF THE STATE Grouse, like other wildlife, are the properly oi all the people. Individuals and organiza- tions, therefore, rightfuU) look to the State for leadership in planning and carrying out any overall projects in behalf of the bird. Here the State is confronted with a difficulty of long standing. Naturally, concern for and interest in the bird's welfare breeds many a suggestion as to what should be done. The halls of the hot-stove leaguers resound with well-intentioned, vigorous debate. Come hunting sea- son, the State's Conservation Department is deluged with suggestions, complaints, demands, constructive ideas and crackpot impressions. Bedeviled bv such a deluge it is small wonder that many state game administrators are apt to overlook the broad picture in trying to satisfy the immediate demand. In this respect New York is fortunate in that the pertinent facts are already at hand as a guide for constructive action. What, then, ought we to expect of the State by way of practical assistance in producing more grouse? The suggestions that follow fall broadly into two divisions. The first concerns responsibilities of an annual nature, the carrying out of which immediately affect both grouse and hunter. The second group are long-time comprehensive projects calculated to produce a steady improvement in conditions for the bird over a period of years. Annual responsibilities include: — 1. A yearly inventory of conditions and abundance. 2. The regulation of hunting. Long-term projects encompass: — 1. Research into the life history* and management of grouse. 2. Application of the facts thus found. 3. Acquisition and development of public lands on which grouse represent one of the major crops to be encouraged. 4. Maintenance of adequate hunting opportunities. 5. Encouragement of private landowners to produce more and better grouse coverts. 6. Stimulation of an interest in and a knowledge of grouse problems, and their solution. 7. Sustained efforts on fundamental game, soil and forest conservation programs. To carry out any one of these is a job in itself. Attendance to all of them, even over a * Practically complete for New York. 602 MANAGISG GROUSE AREAS period of years, requires an organization and funds beyond tiie reach of many states, partic- ularly since some ])robiems are more pressing than others. .New York is among the fortunate in that the interest, the funds, and the techiiiial assistance prerequisite to attacking all of these simultaneously have already been put to work on these problems. With a background of experience thus gained, let us see what part a state can pla\ in improving conditions for grouse. Annual Responsibilities Yearly Inventory Every business takes a periodic inventory of stock and conditions. Faced with a species that fluctuates widely in abundance and an ever-increasing hunting pressure, this, for grouse, is an absolute prerequisite to any intelligent adjustment of season, bag and take limits, to sup- ply and demand. It may be secured in several ways. These are described in Chapter XVII beginning on p. 675. Regulation of Hunting Hunting, with grouse, is not the hele noir it is generally supposed to be, but, where sur- pluses exist or populations are scarce, the necessity for reasonable control is clearly recog- nized. So also is the State's responsibility for making its regulations a reflection of the con- dition of the grouse crop. Without a good annual census this is impossible. Since the fall crop is dependent, not alone upon the number of birds in the winter coverts, but also upon nesting and brood success throughout the succeeding spring and summer, it is obvious that such an inventory cannot be completed before late summer. For this reason a state legis- lature, in session the previous winter, cannot possibly have the facts at hand with which to intelligently establish open seasons and bag limitations on the daily take. Realizing this, New York has wisely placed the responsibility for setting such regulations on the Con- servation Commissioner who is in the best possible position to decide on the yearly restric- tions in accordance with conditions. The law transferring this responsibility requires that the regulations be promulgated not later than September 15, and wide publicity is given to them thereafter. In this way alone can the season be made to best serve both the grouse and the hunter. As the needs of the situation are more clearly understood, other states are grad- ually adopting a similar course. Long-term Projects Research Facts are the fabric (lut of which any ]iatt(rn of successful management is woven. Rut research is at best a slow and costly undertaking, with results seldom directly translatable into dollars. Public agencies, largely federal and state, therefore must accept the lion's share of the work of carrying out factual C-\i)lorations. When* funds are insufficient to meet state needs in this respect, there still exists ihc o|)|ioitiinil\ for collaborative work with a university. A few thousand dollars thus spent ma\ bring substantial returns. The Investi- gation is an exami)lc of what ma\ be aiconiplished In enlisting assistance from many quar- ters. In some cases it will be possible to adapt basic studies carried on elsewhere to local conditions. THE ROLE OF THE STATE 603 In all but a few states the necessity of employing men technically trained for such work is recognized. Often, they may serve a dual purpose by acting as consultants on wildlife prob- lems in addition to carrying out their research. Service to the Landowner Facts are largely valuable in proportion to their use. Realizing this, the Conservation Department has divided New York State into eight game management districts. In each a trained technician is available to advise on grouse and other wildlife problems. But assisting landowners and answering questions is only a part of their assignment. They are also respon- sible for purchasing State game lands, planning productive developments thereon, and see- ing that they are carried out. If the State is to fulfill its responsibility in these respects, some such an organization is a prime prerequisite. Acquisition ami Development of (Arouse Lands In New York State it is no longer considered necessary to acquire public lands to serve as refuges for grouse. Emphasis instead is placed on |)roviding public hunting grounds and NEW YORK STATE .S CAME MANAGEMENT AREAS ARE DEVELOPED ON THE MULTIPLE USE PRINCIPLE on managing the cover thereon so as to produce as nearly maximum crops as possible. On public lands where grouse are not the major crop it is usually possible to make conditions more favorable for them by suggesting, to those in charge, practices and improvements which may be carried out incidental to the major development of the area. Much can be done in this manner to encourage better grouse habitats on state lands where recreation or forestry are the main considerations. C)U1 MANAGING GROUSE AREAS Maintaining Hunting Grounds In New York, where there is at least one grouse hunter for every 50 acres of grouse cover, it is obvious that the State can never be expected to own sufficient public hunting grounds to supply adequate hunting lands. Only in the vicinity of the larger centers of population, there- fore, is the State justified in purchasing and developing aii) substantial areas primarily for this purpose. Even this is dependent upon the availability of low-cost lands and their sub- sequent management to provide recreation and a forest as well as a wildlife crop. This does not preclude the desirability, however, of acquiring large areas of abandoned land in other parts of the State for such purposes. The principle of State purchase and restoration of such lands to productive use is a sound one. Properly directed, such a project cannot but sub- stantially increase the amount of land open to public hunting. Because the ruffed grouse is seldom found in heavily farmed areas, less cover is posted against grouse hunting than is true with many other species. W here posting becomes serious, the State should consider remedial measures. In this connection the establishment of land- owner-sportsman cooperatives, such as have been developed to meet similar situations in pheasant areas, might be tried out providing the grouse are sufficiently numerous to warrant the expense. Stimulating Grouse Production on Private Lands Though obviously desirable, few states have made even a start in stimulating private land- owners to produce more grouse. Yet, until ways of doing this are worked out. the State can exert but little influence on the majority of grouse covers lying within its borders. The dis- trict manager setup already mentioned stands ready to advise and help those landowners who have the interest of the bird at heart. To reach the others, it will be necessary to provide some other quid pro quo to stimulate constructive action. Stiniulaling Interest and Understanding Another contribution of the State lies in stimulating an interest in and an understanding of the grouse problem. Books such as this, articles, talks, discussions and movies all help. So also does the formation, with State encouragement, of local conservation clubs. Only by starting an educational program with the youngsters in the schools, however, can we. in the long run, get across the story of the value and problems of wildlife in general, and of grouse in particular. The school lecture program of New York's newly created Division of Conser- vation Education represents a long step in the right direction. Emphasis on Fundamental Conservation Programs By now it is clear that whatever helps coiiserxalion in general, sooner or later is likely also to benefit grouse. Basic farm and forest, soil and wildlife conservation projects should be vig- orously supported after making certain that adequate consideration is given therein to improv- ing game conditions. Reading this chapter, it may seem to some that any effort they can make to better grouse conditions must be so small as to make it unimportant. Nothing could be further from the truth. Granting the enthusiasm of the authors for their task, the material for this book could never have been devoloix'd without the liel|) and backing of thousands of grouse-con- scious individuals. Oaks grow from acorns. Inherent in the germination of every seed of interest and activity lies the probability of making conditions better for this grand game bird. CHAPTER XV DESIGNING GROUSE COVERTS AND SETTING UP MANAGEMENT PLANS* By Gardiner Bump THE ESSENTIALS OF GOOD COVER DESIGN Integration With Other Uses COVER ORGANIZATION Planting Design for Open Land — Overgrown Land Design — Woodland Design — Clear-cut Units — Forest Stand and Game Cover Improvement PLANNING COVER IMPROVEMENT Establishing Management Objectives — Determining Current Cover Conditions — The Cover Survey — Cover Type Mapping — Analyzing the Covert in Terms of Grouse Needs — Recognizing Cover Deficiencies — Locating Hunting Areas and Seed Stock Refuges ORGANIZING THE OTHER FORCES OF PRODUCTION Predation — Disease — Control of the Harvest — Dispersion of the Birds — Some Generally Unproductive Practices — Providing Artificial Food and Shelter — Restocking SETTING UP MANAGEMENT PLANS Basis — Organization of the Plan — Background — ^Objectives — Division of the Man- agement Area — Covert Surveys — Analysis of Cover Conditions — Development and Maintenance Plans — Regulation of the Harvest — Coordination with Other Uses — Protection and Other Special Problems — Appendix. ^ SUMMARY Nature produces the plants and animals that make up the environment in which the grouse lives. Left to her own devices, the quality of the coverts produced, influenced as it is by a variety of conditions, may be good or poor. It is perfectly practical, however, for the sportsman, landowner or wildlife manager to plan and carry out changes in these to the end that better coverts will result. These improvements should be made in ac- * The word "coven" is here used to represent the vegetative cover and physiographic features within a more or less distinct unit, separated from other units by some easily recognized features such as open lands, roads or water. A covert may contain one or many grouse habitats plus some relatively unproductive areas. f)()6 DESIGM.\C GROUSE COVERTS A.XD SETTING UP MANAGEMENT PLANS cordance with a definite plan. Such planning is here called "design." (p. 607). A clear understanding of the essentials of good cover design will be of material assistance in determining what changes are desirable and how they may best be made. I p. 607). The basic component of grouse coverts is the cover type. Both fundamental and incidental needs of the grouse are satisfied, but in varying degrees, according to season by these types. In planning cover improvements, the i)urpose each serves must be kept constantly in mind. (p. 608). The arrangement of cover types in large measure controls the indductiveness of the indi- vidual habitat as well as the number of habitats to be foinid in the covert, (p. 609). The possible combinations of cover types that collectively make up a grouse habitat are legion. Ten of the simpler, most conmion ones are diagrammed and analvzed bv way of illustrating their \alue for grouse, (p. 610). Since the greater the number of good habitats in a covert, the larger the number of grouse likely to live there, the arrangement of cover types to produce the maximum number of habitats practical is discussed and diagrammed, (p. 612-613). It is quite practical to produce subsidiary crops such as forest products and recreational op- portunities, as well as to assist in controlling soil erosion and balancing water runoff, along with the encouragement of a good crop of grouse. Consideration should therefore be given to the integration of these other uses in planning grouse habitats, (p. 615). Productive patterns for open lands (p. 616), overgrown lands (p. 618), woodlands (p. 620) and clear-cut units (p. 621 ) are here outlined, to give an idea of the changes which one may undertake by way of making these types more adequately fulfill the critical require- ments of the grouse. A survey of existing conditions and influences, both plant and animal, that may affect grouse, is a step prerequisite to the organization of a]j]>ro|)riate development plans. The inten- sity with which this .should be carried out is dependent upon the extent of the improve- ments contemplated, (p. 626). A cover type map is an almost indispensable aid in effecti\ely anal\/inj.' llie covert in terms of grouse needs, in planning irninovemenls. and in keejiing a rerord of progress, (p. 626). With a good covert survey and cover t\])e map it is easv to recognize existing cover deficien- cies and to locate areas best suited for development as hunting grounds or refuge units, (p. 627). It is sometimes necessary also to conlrol. In sonic extent, the other forces inflncTicing produc- tion such as |)redation. disease and Ininting. and to restock dcplclcd areas. Kach of these must be considered in an\ well ordcn-ij plan. I p. 629). Management plans for developing and maintaiiiiiij; a grouse area sIkuiM be ^cl up in incliulc certain definite objectives. Since the items which should he inchuled in sucli a plan have not. as yet. been standardized, a suggested organization of these is here described, (p. 632). THE ESSENTIALS OF GOOD COVER DESIGN 607 The preceding chapter painted with a broad brush the picture of what may be done and what to avoid in improving conditions for grouse. In this chapter are pencil-pointed the specific procedures and practices by which covert conditions and needs may be analyzed and the forces controlling production marshalled in accordance with a pre-determined plan of management. Succeeding chapters offers suggestions as to how cover and crop, predators and buffer species, may be so handled as to maintain continued productivity. These are presented in considerable detail to provide as broad a basis as possible for those interested in the intensive development of grouse coverts under the variety of conditioiis characteristic of the bird's extensive range. No one realizes more clearly than the authors that grouse management is scarcely out of its swaddling clothes. Under such conditions the procedures here suggested must not be accepted as the only or perhaps even the best ones likely to accomplish the desired results. They are suggestions to be modified as experience directs rather than cut and dried prac- tices. But they are tried and workable. As such they represent a concrete starting point for those who would venture into the relatively uncharted field of making two grouse live tomor- row where but one exists today. How then does one go about analyzing and organizing grouse coverts according to a busi- nesslike plan? Architects and land-use ex])erts call such |)laiuiing "design", though wildlife managers have not yet widely adopted the term. The architect plans a structure to meet a given use; the manager designs a game covert to produce game and other crops. Woods, fields and fence corners, rather than steel, lumber and stone, are the materials with which one must work but the fundamental steps are the same. In either case satisfactory results are dependent on good design. THE ESSENTIALS OF GOOD COVER DESIGN The principles of cover makeup are presented in Chapter 111. Covert and type size and shape, composition and arrangement are discussed on pages 111 to 11. '1 The role played by the latter points is so decisive as to warrant a s-eparate section, found on pages 168 to 170. The plant species, important as food, are detailed in Chapter IV and the makeup of ideal coverts has just been pictured in the previous chapter. While these discussions provide the background for understanding effective cover organiza- tion, a few points may well be amplified. The first of these concerns the role each cover type plays in fulfilling grouse cover needs. Grouse are adaptable birds. No one of the many types of cover, that make up their hab- itat, fails to provide some food and shelter at one or another season of the year. Since much of this is, however, incidental, table 94 has been prepared to illustrate the principal require- ments fulfilled by each type. Lest confusion arise between this and statements made in Chapter III. one must have clearly in mind that this analysis is based not as much on the use made of each type as on the extent to which they meet the major food and shelter requisites at the time in which they are impor- tant. For example, patches of conifers are most frequented during eight of the twelve months of the year. Yet only in providing winter shelter do they supply a prime need, not also adequately filled by one or more other types. Thus, in winter they are of primary im- portance. As another case to point, in spring many adults frequent mature woodlands where available. Yet several other types are also largely used for feeding and resting. But the basic f.08 DESIGNING GROl'SE COVERTS AND SETTING UP MANAGEMENT PLANS Z ■< H CC O a. "-; a J E ^^ SS u s ii QX 3 z W a. >- H a; > z Zh ^ = H 3 3 b Q Z < < a '73 ? CS Z Ed H -a 3 C^ o o e> a.c.u a Soi a a o ii o i - o *- « M w •- < : o c 2 2 P. 73 « O V o o o o -3 5 O O 0 o ^ o 2— 2 o 2= On o o o U d -J C. £0 S. u o a H" 5 o «> - 5 -s S = P J a; cc !T3 ■§ -2 12 -S = — c — = 0 3 - t t**— r i^ suQ ■am EI.U.US »>, ^E3"i' 5-2 I I'l ii l'**a'i'*S-c 3 1 X.O E S c X ■2* ¥ 5 » -♦-♦-♦••••wo 9 i THE ESSENTIALS OF GOOD COVER DESIGN 609 spring activity is reproduction. For nesting, female grouse prefer second-growth hardwoods. Thus, at this season, this type occupies a position of special importance although, in actu- ality, it is less used for other purposes than are mature hardwoods. In other words, many types may provide food and shelter for the incidental activities of life but only a few seem adequately to meet the major seasonal needs that must be provided for if the bird population is to thrive. A second point, always in need of emphasis, is the importance of proper cover arrangement. A thousand-acre tract of dense woodland usually attracts but few birds except along the edges where it adjoins overgrown fields or slashings. Broken up into smaller cover units, an area of the same size will be used to a much greater degree. Likewise, hedgerows intersecting open fields may furnish excellent fall food but will seldom be used by grouse unless the other components that collectively make up good grouse cover are close by. The arrangement of cover types in large measure controls the productiveness of the indi- vidual habitat as well as the number of habitats to be found in the covert. Let us consider each of these points separately. To visualize the part cover arrangement jjlays in producing the individual habitat ten of the simpler combinations, commonly found in the Northeast, are reproduced in Figure 63. Alongside each is an analysis of the extent to which the combination meets the four basic grouse requirements as well as the probable result in terms of grouse habitat. In presenting these, it is of course recognized that any one component, such as "over- grown lands" may in actuality be made up i.i«. A txiTjil Ihnl fairly l-)iimli(-rc(I hariluiMHU nii«] conifpr* occasionally may niiitaiit a liuHit'ii'nt variety i.f vi-grtalion ^o as to meet all four jirimary nrt-ds. In years of ImbI) b'o""*' populations some birds arc usually l<> hr found in siirh situations, t If ronifioscd of birch, pofiplr, or aider patrlies. j If overKiown lands are composed of brusli cliaracloristic of old pastures, abandoned fence corners, etc. S Conifers is the only type in which winter feed is deficient. This forces the birds to seek food in tile adjacent ovcrsrown lands in which escape co%'er (from predators) in winter is usually deficient. THE ESSENTIALS OF GOOD COVER DESIGN 611 Usual Habitat Combinations (Continued) Winter shelter Spring breeding grounds Summer feeding grounds Fall feeding grounds Fair habitat for grouse Analysis — good — excellent — good — poor Winter shelter — poor to fair Spring breeding grounds — excellent Summer feeding grounds — good to excellent Fall feeding grounds — good to excellent I'dor to fair habitat for grouse Winter shelter Sjiring breeding grounds Summer feeding grounds Fall feeding grounds (idiid habitat for prouse Winter shelter S|ji ing breeding grounds Sununer feeding grounds Fall feeding grounds GoikI habitat for grouse ■good to excellent • good ■ good to excellent good to excellent ■ excellent good to excellent • good to excellent fair^ to good* Winter shelter — excellent Spring breeding grounds — good to excellent Summer feeding grounds — good to excellent Fall feeding grounds — good to excellent I Excellent habitat for grouse KEY TO FIGURE 63 OPEN LAND ^j>^ HARDWOODS a CONIFERS I ■ ■ ■ [:;;;:j overgrown land KvI3 conifers P^^^ SLASHINGS WM\ hardwoods 612 DESIGNING GROUSE COVERTS AND SETTING LP MANAGEMENT PLANS to seek buds and berries in the adjacent overgrown lands. Here escape cover from predators is often deficient in winter. This is one of the reasons why from 10 to 20 per cent of a pre- dominantly coniferous plantation may well be planted or left to grow up to food-producing broad-leaved species. The more such hardwoods can be scattered through the plantation, the better are the conditions for grouse. Another common grouping (6) is to have conifers surrounded by hardwoods. Conditions here are likewise unfavorable for grouse because of a lack of productive summer and fall feeding grounds. The same may be said of hardwoods bordering mixed hardwoods and conifers (combina- tion 7) since no provision for variety in summer and fall feeding grounds, as represented bv slashings and overgrown lands, is made. The above cover groupings are often associated with regions where the land is being, or was once, farmed. In the more heavily wooded regions of the State, few grouse are com- monly to be found except where man or nature has made substantial breaks in the forest cover. Such situations often result from lumbering or from land clearance followed by moderate pasturing. Where the cut-over areas or slashings are surrounded by hardwoods (combination 7), adequate winter shelter is lacking in the latter t\pe and is provided but moderately in the slashings. The result, in terms of birds is, accordingly, disappointing. On the other hand, where the cover surrounding the slashing is composed of mixed hardwoods and conifers, as in combination 8. this deficiency is eliminated. Providing a slashing is old enough to furnish excellent summer and fall feeding grounds, the result is usually a rather productive habitat. Perhaps the best combination fairly frequently encountered is where mature hardwoods and conifers are to be found adjacent to second-growth, which is in turn adjacent to a belt of overgrown fields (combination 9). Open up the cover a bit more by making a few small slashings in the hardwoods and conifers and the habitat becomes as nearly perfect as arrange- ment of cover types can make it. This is illustrated in combination 10. From the al)o\e it is evident thai type arrangement, as well as coin|)(isition. plays a decisive role in determining the number of grouse an indi\ idual liabitat can produce. It is equally true that the greater the number of good habitats in a c()\ert. the larger the nund)er of grouse likely to live there. Conversely it is ])erfectl\ ])ossiblc to have an adequate aninunt of each cover type present in a given covert and still produce hut few grouse. This ])rinciple is illus- trated in figure 64 where both areas contain the same amount of each important cover type. Yet onlv in "B", where all types are within short distance of each other, does more than one really productive grouse habitat exist. Here, though the total amount of each type is the same as in "A", they are so arranged as to furnish many more units of complete bird or brood cover within the same area. To translate this theoretical conceijlion into a practical situation as it might exist on the ground, let us glance at covert I of figure 6.5. Here the only really desirable habitats for grouse occur at the point where hardwoods, slashings and mixed woodlands meet and along the dashed line between the laller l\\(i Ivpcs. One would exi)ect to find but few grouse in a covert so composed. By way of illustrating the o])posite silualion. coNcrl 2 idntains about the same amount of each type but so arranged as to provide not one Inil clcvi-n piniluctixc hab- itat spots (identified bv circles) as well as seven particularly desirable edges. Tlie normal level of abundance in this covert would exceed that in covert I, in most years, many fold. THE ESSENTIALS OF GOOD COVER DESIGN 613 -.t-f- '• STRING BREttlNO Si'o. b^li:" ; TALL FtEDiwa :'.;:;: !•■•••/ wirms sielter/) ■.••.■■■■.■■■■-■■. ■.■•..■.■-.r..f,'.'I'.'.'.V.'.*.V»ViV«i Nucleus of Grouse Hobitol FIGURE 64. THEORETICAL DISTRIBUTION OF THE SAME AMOUNT OF CRITICAL GROUSE COVER TYPES TO lUASTRATF TIIK IMPORTANrK OF PROPER ARRANGEMENT SCALE I 1000 ^ ^^ k &iMs;u:;;/V:.?:^::;:^:::::::::::::::;>'^ © □ O^RCROWN LAN'D m ". KEY TO FIGURE 65 AROWOCIS ^i^M HARDWOODS 8 COWFERS t"Xl CONIFERS SLASHINGS FIGURE 65. A PRACTICAL ILLUSTRATION OF THE EFFECT OF COVER TYPE ARRANGEMENT IN DE- TERMINING THE NUMBER OF GROUSE HABITATS IN A COVERT r)14 DESICM\'G GROUSE COVERTS ,1X0 SETTING UP MANAGEMENT PLANS In studying the combinations illustrated above, it is of course realized that trees and shrubs, unlike lumber, canriot easily be moved about and put in place to quickly translate a desirable pattern into reality. Much can be done, however, to alter the existing arrangement of cover types in any covert for the better over a period of years, if the principles and ends are well understood. Another major point, entering into good cover design, is the actual composition of each type that collectively comprise the covert. While this has been described in detail in Chap- ter III, it is well to emphasize here that grouse cover is not static by nature. Thus the makeup of a type is constantly changing. Take, for example, a clear-cut area. An eight vear old slashing may furnish excellent, a two year cutting but fair, summer feeding grounds depend- ing upon whether the growth is open or dense and the food-producing species are abundant or scarce. Inherently, however, slashings have the capacity to produce good summer food. Primarily they serve this purpose. Secondarily, they may also afford considerable fall food. As the character of a cut-over area progresses from herbs and berries to predominantly sprout growth, it becomes progressively less attractive to grouse. If growth continues unchecked by cutting, succeeding years will see such a unit change completely in type from a slashing to second-growth forest. Though it still will be frequented to some extent by broods in summer, its main function then will be to ])rovide spring breeding grounds and next, some winter shel- ter if conifers are present. Other types, notably overgrown fields, also may change rapidly in composition. Allowance for this should be made in designing and maintaining coverts. A final major ])oint. not to be overlooked, is the effect type size and shape may have on covert productivity. The size of a cover type determines the amount of space inside an area while its shape controls the amount of edge enclosing that space as well as the greatest dis- tance at which any point within it is situated from the edge. Thus a square or a circle has less edge than a long, narrow strip or one irregularly shaped. Since edges may be an important component of good game cover, it is wise to encourage the longest possible outline while retaining sufficient depth. Just how far one may go in this direction will depend a great deal on the physical condition of the site and the uses to be served. While coniferous reforestation may furnish admirable winter shelter, much of it is wasted, so far as grouse are concerned, if it is deep. Only a strip along the periphery approximately 200 feet wide is likely to be much used. The normal degree of use in most types of cover tends to be inversely proportioncil to the distance from the edge. A small slashing max allract mam birds. As the distance across the cover type incrcas<'s l)e\(ind llic minimum ticcds. utilization ])er unit of area decreases because the liirds have more fnpiii wliicli to choose. They may occasionally travel deeper into cut-over areas or brnshlaiids. at times going half a mile or more from their usual haunts. More than P)0 |nT ccnl of the grouse nests recordc!v!v!*!-;v;v!*i-i-iv!*!\v^ ■'.■.■'.•'.■'.■'.■'.•'.■'.■'.\- ;.»;.;• :'://.■ ,V .*.'.'. •:•:•:■;■:•:•:■:■:■:•:■ ivXvX::::::::::::::::::^^^^^^^^^ *.•.'.•.*. :•:•>:•:• /.•.•/.••.■.•.•.•.•.•.•.•.•.•.•.•.•/.•.-.•,•.•.*.•.•.• ^????Wm^^^m^^??7^^^^^^m^^' .. . .!■■.■.■■■■■.■■■■■■.■ A THEORETICAL [a??v1 Hardwoods B PRACTICAL Conifers FIGURE 66. SIMPLEST DESIGN FOR PLANTING OPEN LAND TO PROVIDE SOME GROUSE COVER J COVER ORGANIZATION 617 .■.".* '. .'. .*. .'.'.'. •:•:•: W--y':- .'.■.• '■'.'. ■'-'■'■'•'•"-■ •/'• '^^^■y^:^- ■■:■■:■■:■■:■■:■■:■ •.•.•.•.•/.•.•.•.•.•.•.•.•.•.•.•■.' m^^^^^^^^ J :.:-v:v:v:v:::::v:^ J -J \"i , rt fl fl n a-Mm •*•* ■••'■•-•| 1 1 1 H 1 . t . ' ■.'.".■.•.•,•.■ ' ■ ■ ■■' .'.'..'.',■.'.'.•.■.< '.'.'..'.'.■.'.'.-.■. \.' .'.'.'.'.'.'.' .' ','.'.'.',','.'.'.'.'. "f .■.■,■.■.'.'.■.',' '.■.■.'.■.■..■.■,".■. :•:•:■:■:■:■:•:•:■:•:■!: •,■•:•:■:•:■:•:•:•:■:■'■ -v-'-'ivi'-v .■.•!■!•:■!•!■!■:■!■!■: ;.•/.•.•.•.•.•.■. ".*.".■,".'.'.'.'. ■'.■-■-'.■.'.'.■.'.■.■ . I'. *.*,',*.' * ■ • '.*.■.*.'.•.•.■.*. •.■.■.■,■.■.■.■.■.-.•. -f •.■.•.•.'.•.'.•,•. ■:-^>y<<-:-::^^^^^:: .■.'.'.'.*.*.'.*.' ''.•.•.•.■.■'.•.■'.•'.■'.•'.■ WW '.':.'.' WM^'-^:'^^ :•:•:•:•:•:■:•:•:• t A THEORETICAL B PRACTICAL I I Open Land Fx??| Hardwoods Shrubs |^%^ Conifers FIGURE 67. MORE PRODUCTIVE DESIGN FOR PLANTING OPEN LAND TO PROVIDE GOOD GROUSE COVER by just two types of cover — mixed woods and overgrown lands. The simplest design, then, which might be followed in planting open land to provide these combinations, would be a series of units as shown in "A" of figure 66. Because of planting costs, though, it might bet- ter be organized as shown in "B". In the most productive coverts analyzed, an advantage seems to accrue when winter shelter and spring breeding grounds are separate and when summer and fall feeding grounds are, likewise, represented by different types. In New York the best winter shelter is furnished by conifers, a block of which is accordingly included in suggesting the design of a theoretically productive grouse habitat (figure 67). A narrow strip of open grassland, from 30 to 50 feet wide, may well separate each group of habitats. The Investigation has been unable to demonstrate any positive value accruing from such strips, yet those coverts which grouse use the most usually contain such openings. These strips can be made into good firebreaks. The best summer feeding grounds over most New York grouse range are clear-cut areas or slashings from three to ten years old. Obviously this type cannot be utilized in designing grouse habitats arising from lands to be planted. Unless woodlands, in which small clear- cut units may be established, are found adjacent to open fields, it is still necessary to com- bine summer and fall feeding grounds in the form of edge development. With the right species properly arranged in strips across open fields ("B" of figure 67), a 6]P, DESICMXG CROrSE COVERTS AND SETTING UP MANAGEMENT PLANS combination may be secured which should produce not only a good crop of grouse but also of other forest products. As regards the size of each type to be planted, no minimum dimensions have been deter- mined. The area required is, in part, dependent upon the composition of the type some years after planting. Naturally this, in turn, is influenced by many local factors. To avoid diffi- culty the dimensions of each type suggested here are considered to be sufficient assurance against possible inadequacies. The open land development pattern common 1) used in .\ew York provides for winter shelter by utilizing a central strip of conifers from 300 to 500 feet in width and of any desired length. Here the trees may well be planted 8x8 feet apart, so that a crop of forest products may result. However, if grouse are the sole consideration, the spacing might better be further apart unless the site is poor. This is less expensive, results in a bushier tree and tends to encourage a more desirable mixture of conifers and hardwoods by giving the latter more open space in which to establish themselves naturall). This band may be flanked on either side by strips from 200 to 300 feet wide in which conifers and hardwoods are planted in irreg- ular clumps to furnish spring breeding grounds. When such an area can be placed close to existing woodlands, it may be practical to plant only the conifers, leaving the hardwoods to seed in by natural means. To allow for fall feeding grounds it is suggested that a strip from 50 to 100 feet wide be left. Over this, at irregular intervals, may be planted small clumps of conifers edged with broad-leaved trees and shrubs that normally produce attractive food at this season of the year. Theoretically these strips might well be placed as indicated in figure 67. although, in actu- ality, there is no value to the geometrical regularity of the design. In fact, a scalloped outline is desirable. Full advantage should be taken of any food species already present. Hedge- rows, woods' edges and stream borders, where attractive plants normally occur, make ideal units around which to organize a fall feeding strip. The small, open strip previously men- tioned may then be left before repeating, in reverse, this planting design. Overgrown Land Design In New York State, Nature is a past master at producing brushy spots attractive to grouse in summer and early fall. Most of these result from the abandonment of once-cleared fields, pastures, or from hedgerows. Some predominant characteristics which make them desirable are: — 1. A variety of trees and shrubs, many of which furnish feed as well as shelter for brood and adult alike. A scattering of such species as apple, thornapple, cherry, wild grape, viburnum and dogwood, throughout overgrown lands materially increases their usefulness not alone to grouse but to Tnam other forms of wildlife a> ucll. 2. The replacement of grass, asters or goldenrod bv other ground covtM- more character- istic of a woods" edge environment. 3. In general, a rather open stand of trees and sliriii)s although the presence of occasional small, dense thickets of shrubs seems imt to be (lelririieiital. 4. Few, rather than many, conifers. Every grouse hunter ])ictures, from his own experience, the necessary composition of over- grown areas for it is there that birds are most likely to be found in the early part of the COVER ORGANIZATION 619 hunting season. Little space need, therefore, be devoted to describing them. In the Northeast, perhaps the most common situation involves the development of grouse cover from an area of abandoned farmlands and woodlots. The former are usually more or less overgrown. Thus is provided an existing pattern on which to build. Figure 68 illus- trates such a development. Advantage should be taken of food and shelter already present whenever possible rather than to attempt the relatively expensive procedure of artificially producing it. Overgrown fields usually furnish good fall feed. If they are large, the parts most strate- gically located and productive of fruits should be maintained as fall feeding grounds. The — LfcCENb — Exis^rinj WooctstBrujhlAnd z S AAA >' FIGURE 68. GROUSE HABITAT DEVELOPMENT ON SUBMARGINAL LAND Existing woods and bnishland (1) Overgrown fields (2) Hardwoods (3) Hardwoods and conifers (4) Conifers (5) Cut-over areas Explanation Open land development (6) Open land — not to be planted (7) Overgrown edges — to be planted to food produc- ing shrubs and trees with occasional clumps of conifers. It may also be left to seed in naturally on the lea side of the woods or hedgerows. (8) Hardwoods and conifers — to be planted or left to seed in naturally according to the situation. (9) Conifers — to be planted tvith mixture of conifer- ous species 620 DESIGNING GROUSE COVERTS AND SETTING UP MANAGEMENT PLANS rest will naturally grow up into spring breeding grounds. One of the toughest problems is to prevent this transition from taking place where productive fall feeding areas are already well established. This may be accomplished by periodically removing the taller saplings that threaten to choke out the other trees, shrubs and ground vegetation on which the birds rely for food. A method of poisoning the less desirable species, to prevent sprouting, is described in the succeeding chapter.* Clearing with an axe is expensive since, normally, but little use can be made of the cuttings. Moderate grazing hy cattle tends to hold back the natural development of brushy areas into forest although patches of unproductive grass may be encouraged thereby. \^'Tiere over- grown fields are pastured adjacent woodlands should be fenced off. Even in case maintenance of the necessary amount of overgrown land proves to be too costly, it is always wise to release from competition any particularly desirable food producers such as apples, thornapples or clumps of dogwoods or viburnums lliat may be present. Conversely, where trees or shrubs of the right species are scarce, as in an abandoned meadow bordering a woods, it may be justifiable to introduce suitable species by planting if they do not appear to be seeding in naturally. This, also, is discussed in the next chap- ter. Woodland Design With but few exceptions, not too much can be done to improve the forested parts of a covert except over a period of years. By selective logging of parts of a woodland, small openings with their attendant shrubs and ground cover may quickly be created. A thinning of crooked or diseased species of low value to game or as timber will, likewise, open up a thick patch of woods. Skillfully planned and carried out, such practices generally tend to improve both food and shelter for grouse. However, care should be used not to encourage too dense an undergrowth as the birds are more partial to a woodland with moderate undercover. Where grass is likely to be encouraged by heavy cutting, this should be avoided. Even light graz- ing, except where undergrowth is dense, is definitely detrimental. In planning woodland improvement, conifers, either scattered through the woods or local- ized in clumps here and there, are particularly to be encouraged since they furnish escape cover and winter shelter. The only exception to this is where second-growth hardwoods, in the vicinity of openings, are available for use as spring nesting grounds. Where winter shelter is desired but not present, it is usually best to plan for coniferous plantings in adjacent fields and in small openings in the woods. Underplanting, unless the crown cover is open, is seldom successful for both root and crown competition in woodlands are usually severe. It is wise to encourage woods' roads and small openings within the forest cover. Con- verselv, extensive areas composed largely of a few species are seldom productive of grouse and >li(mlil In- gradually diversified by selective lumbering or by tliinning the stand unless the crown cover is already sparse. In general, uneven-aged stands are preferable since a greater variety of trees, shrubs and ground cover is likely to be encouraged thereby. Lastly, it should not be forgotten that, while woodlands are the backbone of grouse cover. • Promiiing ezpFrimcnls uiing newly dcvrlnpril plant hormoDct and translocating poiaona to climinatr uoileairable trcea and ahruba are at preaent being carried out. COVER ORGANIZATION 621 there is seldom a good reason why they cannot be so managed as to produce, also, periodic crops of pulpwood, mine props, acid wood, cordwood or timber. Thus one may help to defray the costs of the improvements here described and of forest stand improvement practices as well. Specific suggestions as to how both objectives may be met are given further on in this chapter. Clear-cut Units In extensive forest areas where summer and fall food or winter shelter are deficient, one practical way of improving the situation is to cut a few small slashings here and there. These units may often be worked into the covert design so as to fulfill more than one objective. One small slashing may furnish first a supply of wood and later a rich source of food. Where winter shelter for grouse is notably deficient, the tangle of briers, sprouts and vines ^. '^■. AN OCCASIONAL SMALL SLASHI^G \L\KKS E\TE.\S1\ K W I » H II. \ M).s MOKK ATTKACTIVE TO GROUSE which often springs up on cut-over lands may provide a quick though temporary substitute for the conifers and broad-leaved evergreens that normally meet this need. In planning such areas, the possibility of disposing of the wood may often determine whether the unit is to be re-cut at the end of ten to fifteen years to preserve the characteristic slashing vegetation or whether it will be cut in rotation with several similar units. The latter practice is desirable where a market for the smaller sizes of wood is available. Units to be cut in rotation are usually laid out close by one another so as to provide food and shelter year after year. In figure 69 the three divisions of slashing S 1 would be cut in rotation, while S 2 and S 3 would be recut as soon as sprout growth drove out herbs and berries. Repetition of cutting at intervals over the same unit can be continued as long as grass and sedges do not replace the normal herb and shrub cover. The suggestions here outlined covering the character, size, shape, location and rotation of clear-cut units are still too tentative to be called rules. In connection with the multiple pur- poses to be served by the area as a whole and the individual characteristics of the slashing 622 DESIGNING GROUSE COVERTS AND SETTING LP MANAGEMENT PLANS site and of llie iiiunediately surrounding woodlands, tin- followiiif: points are worth consid- ering:— Number and Size 1. It is seldom necessary to devote more than 10 per cent of the total woodland area to clear-cut units. 2. A large number of small, clear-cut units is preferable to a few large ones. 3. To save maintenance costs, both number and size of units cut should be reduced to the minimum necessary to provide adequate food and cover distribution. Shape 1. Since edges are desirable, wherever practical, clear-cut units should be linear (strip slashings) rather than square or circular in design. 2. Scalloped boundaries may be used to avoid cutting desirable trees and to increase edges. 3. Strip slashings should be bent from a straight line wherever an advantage is to be gained, except where there is a likelihood of their being used as sight lines to facili- tate the taking of a game census when desired. Location 1. It is desirable to locate clear-cut areas from 1200 to 1600 feet apart or from other food patches. 2. For esthetic reasons, clear-cuttings should be located where they will not interfere with the landscape value of the woodland. IS one should be placed close to a much- traveled road. 3. Where possible, they should be established to contact and lie between the maximum number and variety of woodland cover types. 4. Sites to be clear-cut should be chosen, when practicable, where the second-growth or mature timber is of minor potential value. 5. To discourage erosion, they should occupy flat or gentle slopes as against steep ones. 6. Clear-cul units lying along, rather than across, the contours are preferable. 7. Poorer, rather than richer soils, should be favored in locating slashings. 8. Sites containing both damp and dry situations should be chosen where available. 9. It is wise to locate strip slashings with relation to possible use as lines of communica- tion or as fire lines in case of need. Cutting Cycle 1. In New York slashings are usually in need of reculting every hn id liflccii years if shrub and herb vcg<'lalion is to be maintained. The lime ela|)sing between cuts varies with the fertility and moisture conditions of the site. The moister, richer lands have the shorter cutting cycle. 2. Where single units are part of a group to be cul in rotation, tlic niuiilici of units present and the Iciiplh of lime necessary to produce salable wood tiiust also be con- sidered in setting the culling cycle. I COVER ORGANIZATION 623 Forest Stand and Game Cover Improvement For the sake of clarity, many of the suggestions made so far have been presented as though grouse were the most important crop to be considered. In most cases there is much to be gained by cropping not only the game but the forest as well. This is the principle of maxi- mum or multiple land use. In creating environments favorable to the production of both forest and game crops, it should be remembered that each has certain basic requirements. Each crop prospers accord- ing to the degree to which these are met. An important difference arises, however, from the fact that the forest itself is the crop whereas wildlife merely occupies the woodland as a habitat. Thus when the wildlife manager, to better grouse conditions, suggests alterations in the forest cover, he may, in effect, be asking the forester to destroy or forego a small part of his crop that game may prosper. It is equally logical that wildlife be controlled when, as with deer, it seriously damages forest reproduction. Failure to realize these simple truths has led to the building up of antagonisms, largely unwarranted, to management suggestions promulgated by either group and to a tendency to discount the importance in the minds of each of the necessity of meeting those require- ments considered basic by the other fellow. The first step, therefore, in integrating the pro- duction of forest and game crops, is to develop a knowledge of and appreciation for the point of view and requirements of each. Once this is done, the problem is half solved. It is desirable, before management plans are made, to establish the order of importance of the various uses to which a given area is to be put. On this decision depends the degree to which any of the suggestions presented here may be applied. If it is decided to give equal enqihasis to timber and to wildlife i)roduction. conflicts in management practices may often be avoided if portions of the woodlands particularly suited to the production of a forest crop are set aside for this primary purpose. Established plan- tations or patches of woodlands of high comnu-rcial value are examples of this point. Like- wise, throughout the area, an adequate amount of certain cover types may be set aside as of primary importance for grouse production. Among these are overgrown fields, fencerows, orchards, scattered clumps of conifers, and small slashings or other openings in the forest cover. Having located each of these primary-use units, the practices necessary to produce the main crops thereon then may be carried out, together with such other cultural activities as may assist in the production of the subsidiary crop. Such an arrangement, clearly under- stood and sympathetically carried out, will help to avoid many of the difficulties charac- teristic of forest and game cooperatives. One of the more difficult decisions to be reached, before any cutting is done, is to define crop trees, wolf trees that crowd out more desirable individuals, and weed species. For New York, crop trees, from the point of view of grouse, are such food-producers as cherry, birch, beech, thornapple and the hornbeams, and such shelter-producing species as hemlock, pine and spruce. Some of these also represent forest crop trees. A conflict of interests is most likely to occur in deciding which are weed species. The classification of these is dependent upon use which, in turn, is based on whether or not better species are present. Thus, where spe- cies furnishing excellent grouse food and shelter are abundant, the kinds normally less at- tractive may be considered weeds unless they are particularly valuable for other purposes such as for lumber. Were the better species absent, however, the latter would then become crop 624 DESIGNING GROUSE COVERTS AND SETTING UP MANAGEMENT PLANS trees. On the other hand, to the forester, thoniapples ami lidinbeains are unimportant; pop- ple, birch and beech are of particular value largely where a local market for such species exists. The degree to which any species fits into a particular situation, then, must determine whether it is to be classified as a crop or weed tree. The designation of individual trees as wolf trees depends upon their classification as a crop or a weed species and upon the extent to which they are crowding out more desirable trees or shrubs. A maple over-topping a thornapple may be considered a wolf tree if the latter species is scarce and a grouse crop is particularly desired. Conversely, if thornapples are abundant and the maple of good shape and important for firewood, timber or for seed, it would be inadvisable to cut it simply to release the tree beneath. A veteran beech overtop- ping other species more used for timber may, nevertheless, be kept to produce beechnuts for grouse. But if it is one of many scattered throughout the forest, it might well be sacrificed in the interests of better timber production. Where the main crop to be benefited by a woods operation is forest products, it is clearer if one refers to the improvement work as forest stand improvement. Vi hen the principal crop is grouse, it is often termed game-cover improvement. Figure 69 illustrates a tract, some of which was originally open land, organized primar- ily for grouse production but adapted to growing a subsidiary timber crop. It will be apparent that good cover organization is a more complex problem than is gen- erally realized. Those who have the interest, time and means to carry out management prac- tices, whether for grouse alone or in combination with other forest products, may be able to make use of many of these suggestions. Others, with lesser means, can still obtain good results by meeting the basic requirements of composition, size and dispersion within the habi- tat, even though man) desirable refinenienls are not possible. PLANNING COVER IMPROVEMENTS With the foregoing picture of cover essentials and organization in mind, there are three steps necessary before one can produce a bluej)rint for cover improvement. First, the ob- jectives of management must be decided upon. Then current cover conditions must be deter- mined. These done, one is then ready to analyze the situation in terms of how well it fulfills grouse needs. Establishing Management Objectives The possibilities of multiple use of grouse areas as well as the integration of forest stand and game cover have alreaily been presented. Obviously the fust move is to decide on the crops to be encouraged. One thinks naturally of firewood and lundier, of places for camp- ing and hiking, of preventing soil erosion and of regulating stream flow as some of the forest attributes which may be combined with tiie i)riinary objective of encouraging a grouse crop. Other game species, such as the elusive deer or the lowly cottontail, may be welcome likewise. Development plans must be modified accordingly. Determining Current Cover Conditions The objectives once outlined, the next step is to take careful stock of present covert con- ditions. The detail with which such an inventory is undertaken depends, naturally, upon the intensity of the contemplated development. This cataloguing of existing conditions as they affect grouse niav be termed the cover survey. If real progress is to be made it is at the same time most helpful to prepare a cover type map from which one can more easily see PLANNING COVER IMPROVEMENTS 625 13 Jo H tj a o o o: IX a u a S Q < 3 O PS o OS > o (J O Z O P ■«: N z <: o e c o u ca o Q. S'3 - • c ;s.E 626 OFSIGMXC GROUSE COVERTS AND SETTING UP MANAGEMENT PLANS the cover improvements that need to be carried out. Later it will form the basis for devel- oping a covert management plan should one so desire. The Cover Survey Through an inventory of the current situation one seeks to learn the distribution of plants producing food and shelter and the various combinations (called cover types) in which they occur. Another purpose is to find out whether these types are so arranged as to produce few or many desirable grouse territories. Incidentallv one may gain some idea of the wild- life relationships existing between game, predator and buffer species and the hunter. The degree of precision to which the analysis should be carried depends upon the pur- pose to be served. If one is merely interested in periodic changes in cover and in the abun- dance of grouse on a given area, a brief reconnaissance will suffice. If time and effort are to be expended on improving habitat conditions, a broad biophysical survey, covering many or all of the items listed below, is desirable. If one is not certain as to the degree of use to be made of a cover survey, it is wise to secure the maximum amount of information which time and funds will permit. Among the items one may wish to consider are: — 1. The distribution and character of the physiographic features of the area — topog- raphy, soils, water conditions and climate. 2. The distribution, size, composition and density of cover types (crown cover, un- dergrowth and ground cover).* 3. The degree to which each type fulfills seasonal grouse requirements, particularly as regards food and shelter. 4. The distribution and relative abundance of grouse predators and l)uffer species. Cover Type Mapping One effective way of securing much of this information is to make a cover-type map of each covert.^ Such a nmp conveys graphically much useful information. With it one may visu- alize the arrangcMicnt of cover and thus more easilv plot the changes which need to be made lliercin. Also, the close ins[)ection which such work necessitates luiturally draws at- ti'iilioii Id many details otherwise likely to be overlooked. The field notes may be expanded to cover as inan\ other |)oiiits. in addition to those here listed, as seems desirable. It is wise to make u() the type map rea^onabh accurate. Where substantial iinpro\onicnts are planned, it is almost essential. Suitable mapping i)ra(tices. in general, follow those described in standard texts on forest management. The technique developed by ihe Investigation is de- scribed in the Appendix^. The scale should be the smallest that will dearly show the requisite details. Tlie Investigation's survev areas were niap|K'(l on a scale of one inch to 10 chains (660 feet). This was found not only to meet the alxivc rcfpiirenienl Iml In facilitate easy acreage computations. Wlierevcr it is possible to sen the map. This may be done by drawing lines from 600 to 800 feet around and parallel to the edges of existing feeding grounds. The rest of the woodlands outside these lines constitutes a zone in which summer and fall feeding grounds are inadequately represented. To remedy this defect, the location of clear- cut units or slashings may be plotted inside the inadequate zone from 1200 to 1600 feet apart and of a similar distance from existing feeding grounds. Thus roughly located, the clear-cut units may be placed more exactly in accordance with the points heretofore mentioned.^ It is less difficult lo determine \\+iere winter shelter or spring breeding grounds should be located. If necessary, the same technique may be applied. The zone of adequacy is prob- ably about the same for the first, but for the second it is represented by a strip seldom more than 200 feet wide along woods' roads, field borders, small openings in the woods or similar edges. No mention has been made of what to do with open lands such as fields or meadows. These may be left to seed in naturally, or, more profitably, may be planted to provide both a game • Sto p. 698. A Sea p. 622. ORGANIZING THE OTHER FORCES OF PRODUCTION 629 and a timber crop at some future date. Detailed suggestions covering this point have been presented. The need for and the positions of adequate food and cover areas thus having been deter- mined, the problem of locating refuge and shooting units still remains to be considered to make the blueprint for improvement complete. Locating Hunting Areas and Seed Stock Refuges High grouse populations are seldom maintained for long in coverts where the surplus is not harvested by hunting. The role and control of the hunter is discussed in Chapter XVII. p. 678 to 679. Well-planned grouse management areas should be laid out so as to facilitate the harvest of the grouse crop. Not that the hunter is given all the breaks — the birds see to that for grouse are famous shot-dodgers. Cover types may, therefore, be arranged occasionally so as to provide many possible shooting opportunities before the birds reach the comparative safety of thick cover. This possesses the double advantage of helping to prevent surplus crops from going unharvested and allowing the less skillful hunter a better opportunity to share in the birds taken, thus spreading the enjoyment over a larger group. Specific suggestions for setting up areas for hunting grouse, are detailed in Chapter XIV on p. 594 to 596. As regards seed-stock refuges, they are of little value where hunting is strictly regulated as on a preserve. Likewise, where dense stands of conifers or hardwoods present a considerable proportion of impenetrable thickets, no artificially established seed stock refuges are necessary as the birds soon learn to make good use of these as escape cover. On the other hand, where grouse coverts are likely to be subjected to heavy and long- continued hunting pressure, or where they are presently so poor as to support but few birds, an occasional small refuge unit, strategically placed, provides some assurance that a fair num- ber of breeders will remain in the coverts at the conclusion of the season. The proper loca- tions for these may be determined by examining the cover-type map in conjunction with field observations. Fall feeding grounds, together with adjacent thick cover, make excellent refuge units. It is desirable to set up the refuge in the midst of good grouse cover so as to provide an abundance of favorable locations nearby for the birds to occupy in the event such coverts are overshot. Assuming a refuge is desirable, the most appropriate size is from 25 to 100 acres. So long as the boundaries are clearly marked, the better the fall feeding and escape cover con- tained therein, the smaller the unit should be. Not over one-fourth of the total acreage of any covert need normally be used for this purpose. The determination of size and location must be based to a considerable extent upon ex- pediency. Easily recognized boundaries should be used wherever possible. Woodland borders, roads and other natural features serve this purpose excellently. By so doing, trespass thereon, as well as the cost of establishment and maintenance will be cut to the minimum. ORGANIZING THE OTHER FORCES OF PRODUCTION While the main emphasis, heretofore, has rightly been placed on designing the best possible food and shelter combinations, it would be in error to assume that these alone are important. The role played by each of the other forces controlling grouse abundance has been described 630 DESIGNING GROUSE COVERTS AND SETTING UP MANAGEMENT PLANS in detail in preceding chapters. A practical analysis of just how each fits into the scheme of grouse existence may be found in the introduction to the Management section of the book.* One cannot read these without realizing that, while each affects the grouse crop in its own way, few are susceptible to direct control. Indirectly, by building better cover and by con- trolling the harvest, the effect of those factors which encourage increase can be strength- ened; those limiting grouse numbers may be made less destructive. Beyond this, as a practical proposition, there is not a great deal that can be done to help the bird except at prohibi- tive cost and effort. In considering the possibilities, one remembers the losses from predation and disease, over- shooting and dispersion, and is apt to wonder if some control measures for these might not be incorporated in any plan for managing grouse coverts. Such items as the providing of artificial food and shelter and restocking depleted coverts are also occasionally advocated. Let us, then, see to what extent these fit into the overall design. Predation Predation, as a force, may work both for and against more grouse, as already described. When grouse are abundant, forcing some to occupy poorer habitats, these birds are natur- ally more likely to be caught by predators. One of the best ways to avoid such losses is to improve habitat quality. Most important in this connection is the arrangement of feeding and escape areas adjacent to each other. Predator control is one of several activities collectively responsible for maintaining large grouse populations in Scotland and England, where the red grouse is produced in almost unbelievable numbers. It is used hand in hand with the control of certain diseases." the building up of excellent cover, and the complete harvesting of the surplus each year. Though millions have been spent on predator control in this country, no similar result has ever been achieved. Wliere predators are not overly abundant, it is doubtful if their control can be justified. True, over small areas, particularly if thoy are isolated, it is perfectly possible to remove most of the resident ground and winged predators by trapping and hunting. But the cost over a large territory is prohibitixc and the results, as previously desrrijied. are by no means as favorable as has been generally pictured. Nor are bounties a practical way of holding grouse predator numbers in check. Cover improvement over large areas supplemented by trapping or shooting of species particularly destructive to grouse, such as goshawks, great horned owls, foxes and weasels. * See p. r.8fl. A Mainly 8irf>tTKylc>»ii*. mti fminj jn niffrti c'^oni'c. rv.,^ -u r?*^ .'^-'\' ORGANIZING THE OTHER FORCES OF PRODUCTION 631 when abnormally abundant in a particular locality, are the only practical methods by which losses from this factor may be controlled. For the latter the cooperation of local trappers may well be solicited. Disease It has been popularly supposed that little could be done to aid in the control of grouse diseases. Insofar as individual diseases go, this is correct. There are at least three general ways, however, of reducing the incidence and therefore the power of grouse diseases in general. The first is to create as many grouse territories as possible within a covert, thus tending to prevent large concentrations of population. The second is to formulate a plan for reducing these concentrations, should they occur, by harvesting the surplus through lengthening the shooting season. With grouse, it is impractical to redistribute the birds by attempting to trap the surplus. The last is to see that predators are not reduced in num- bers to a point where they cannot he depended on to harvest any surplus not taken by hunters. Though the Investigation knows of no exact proof, it is also a generally accepted prin- ciple that the better the cover the stronger the bird and the less likely it is to become weak- ened and fall prey to disease. Control of the Harvest In designing a grouse area it is important that provision be made for adequate control of the harvest. Even with careful planning and control of hunting, grouse numbers will varv considerably from year to year. Therefore an estimate of the season's crop is necessary.* It is advisable to incorporate in the management plan some means whereby the season may be shortened from the normal period or closed altogether if the birds prove to be unusually scarce. Conversely, if they are found to be overly abundant it is equally desirable that the season be lengthened so as to allow for the harvesting of the unusual surplus rather than leaving it to be removed In disease and predation. Dispersion ok the Birds Grouse are, by habit, relatively asocial birds. This means that in, periods of abundance, many of the young birds may be expected, in fall, to seek out the less populated coverts. By so arranging the cover as to provide more and better habitats in each covert, this tendency may be minimized except in years of peak populations. * Melhotl3 of censiising are prest-nted on p. 676 .C. . c luiliil;il, ulicn |irii|i(il\ i mil j ullcd may be used Id licl|i riii|irove liotli fdud and >licl|ci. (p. (ill). Eire is useful in tliinniniisc cover and setting back tlie succession. It is poten- tially the most dangerous method of all, however, and the cost of control may there- fore be high unless expertly handled, (p. 642). Covert conditions vary with the site and with the past history of the cover. In producing bet- ter grouse habitats, it is up to the individual to choose the method best suited to each situation. The adaptation of each method to the j)roduction of better fall feeding grounds, spring breeding grounds, winter shelter and summer feeding grounds is therefore de- scribed in some detail, (p. 642). Where desired there are a number of ways in which game cover improvement work may be correlated with other forest practices to produce a forest as well as a grouse crop. Where these are combined it is important to select a timber crop, the encouragement and har- vesting of which will result in larger grouse populations, (p. 650). Forest stand improvement work may be carried out to encourage both timber and game production by adapting the practices here described, (p. 651, 777). The control of destructive forest insects and diseases, as currently practiced in New York State, is seldom detrimental either to grouse cover or to the bird itself, (p. 654). The basic principles controlling food and shelter plantings for grouse must be clearly under- stood and closely followed if satisfactory results are to be obtained. Many of the most desirable trees and shrubs are not commonly used in reforestation work and are not particularly easy to establish, (p. 654). The moisture, tolerance to shade and fertility requirements of 32 species adaptable for planting in the Northeast are given in table 97. (p. 655). Generally sjjeaking. hardwood planting stock must be large, with a good root system and carefully planted, preferably by the "center hole" technique to give it the best chance for survival, (p. 660). Direct seeding of nut and mast species is practical only when rodents, such as squir- rels and mice, are not numerous or can be controlled, (p. 661). Conifers seldom develop with sufTicient rapidity to provide good grouse cover when planted under woodland cover, (p. 660). Hardwoods usually require a more fertile soil for successful establishment by artificial plant- ing than do conifers, (p. 661). Shrubs and herbaceous cover can best be established by creating the right conditions and by occasionally assisting natural seeding by judicious planting, (p. 661). White clover, where the soil is neutral or alkaline, may well be encouraged, (p. 664). Better grouse habitats usually result from protecting the cover from fire and grazing though, under certain circumstances, each may be useful in opening up to luxurious cover types or in maintaining openings or brushy cover, (p. -665). ALTERING EXISTING COVER 639 It is one thing to lay the plans to create and iiii]nci\c f;i(ni-,c habitats and quite another to put them into operation. The best set up plans, he lhc> ciini]»li(ated or simple, are of little use unless practical methods of carrying them out arc available. This chapter, then, deals primarily with the ways of improving and maintaining grouse cover. It is not possible to suggest improvement techniques that wiU fit every situation. Some methods, such as those employed in planting open fields to conifers or to certain hardwoods, have been well developed through years of use. Others, for example those designed to en- courage clumps of food-producing shrubs along woods borders, are as yet scarcely out of the experimental stage. The problem is further complicated in that such methods must often be modified to meet individual situations. Here is a condition which places unusual empha- sis on the training, experience and ingenuity of those who are to do the actual work of de- veloping grouse coverts. In planning, one thinks largely of how to make existing conditions better. In carrying out the plans, the practical question of maintenance also demands considerable attention. One has to remember that most grouse cover is constantly changing. Thus nature estab- lishes first one cover type after another, in orderly fashion, until no further change takes place. Fortunately, as previously indicated, these types follow a definite successional pattern. On grouse lands, open fields or grasslands gradually give way to overgrown fields or brush- lands, which in turn grow up into second-growth woodlands and eventually become mature forest. Thus is provided the background against which all grouse management practices must be carried out. The covert management plans may require that existing cover types be maintained, or that they be ahered in order to provide adequate food and shelter in the most productive arrange- ment. To do this one must control succession. It may be set back by removing undesired species or speeded up by means of artificial plantings. There is also the problem of maintain- ing existing types in their present status by retarding or arresting succession. In doing this one is simply trying to bring food and shelter into a more productive balance by changing the composition and arrangement of the cover. In general, the methods by which these ends are accomplished fall into three groups: altering the existing cover, planting new cover, and protection of the existing habitat from forces such as fire and grazing. ALTERING EXISTING COVER* In building up grouse areas, too little attention has usually been paid to the possibilities of securing the desired result by changing the existing cover. The idea of planting the right species in the right place so that Nature may lose no time iji producing the right habitat, some- how has a much stronger appeal to most of us than does the less spectacular proposition of encouraging changes in the existing vegetation. It is only through experience that one learns of the substantial difficulties lying in the path of establishing good cover by planting. Then the desirability of making the most of the existing cover is fully appreciated. Plant- ing has its place, as we shall presently see, but where possible, working with the vegetation already present is by far the surer and quicker way of producing a larger grouse crop. Methods Available Every grouse hunter probably remembers parts of his favorite coverts that, in his mind, * By GardiDer Bump. -~--^^ 'i>'<^i', 640 IMPROVING AND MAINTAINING GROUSE COVERTS are ideal for grouse and others that are not. Li the latter case the difficulty can usually be ascribed to poor composition, to an over-dense cover, or to the size, shape or general arrange- ment of the ty|)es. The cure involves changing the cover. To accomplish this, any one or a combination of five methods may be utilized. They are cutting, girdling, poisoning, grazing and fire. Cutting The usual method of harvesting timber products or of eliminating unwanted trees, is to cut them. \^1here grouse cover improvement can be dovetailed with forest operations, this practice should be followed because of the necessity of salvaging the timber which is left uncut in applying any of the other methods. Cutting is the most expensive, as well as the most certain, of all the possibilities here suggested. However, that many hardwood stumps sprout, following the cutting of the tree, is often a drawback especially in maintenance work. Girdling With species that sprout, girdling ofttimes provides a useful process for checking sucker growth. Care must be exercised to thoroughly separate the bark by a wide cut. Individual trees often take several years to die even though well girdled. In both this and the next method, the trees are left standing, thus providing a likely source of future difficulty. Poisoning Developed some years ago, this practice has been given extensive trials in improving cover on New York game management areas. The idea is to make a wide band of cuts through the bark overlapping each other and encircling the trunk. Into tliese is injected a poison that will kill the tree or shrub and inhibit subsequent sprouting. Sodium chlorate, sodium arsenite, heavy diesel oil, and other plant poisons developed for control work at Cornell University and by the U. S. Forest Service, are used. Though, in the earlier tests, the cuts were made with an axe, it proved more convenient to adopt a poi- soning tool made of hollow pipe, capped at one end and ground to a cutting edge at the other. The liquid is placed within the pipe and, through a valve at the lower end, is fed to the cutting edge each time the tool is vigorously jabbed into the bark of the tree to be killed. The poison is allowed to run into the wound thus formed, thereby putting it in direct contact with the cambium layer which carries the sap. Sodium arsenite was found to be the most effective of the poisons tested. Its major advan- tage lies in its comparative cheapness and its tendency to inhibit sprouting to a greater degree than does girdling. The success of the treatment depends upon the time of year, the species, and the relative vigor of the individual tree. The following table indicates the degree of susceptibility of the species tested. }AJi^m TABLE 95. RFiLATINli SU.SCEPTIRITJTY OF VARIOUS TREES AND SHHUHS 1() I'OISOMNCi WITH SODIUM ARSENITE Sust^eptililn Inlcrmodiutc RcsisUut Ilrmlock PupiT hircli ShailhuHli Fire clii-rry HInck chrrry Wliilo uuti Asncn Yellow birch ItwTh lira oak fJray birch Soft maple llurd niuplc ALTERING EXISTING COVER 641 The disadvantages are that it has been used successfully only on certain species, and then only in the late summer and early fall. When tried in the spring and early summer the com- pound has been known to attract and poison deer which have been killed by the arsenite. As with many newly developed techniques, the results are apt to be uncertain. The solution is also potentially dangerous to handle and the poisoning tool awkward to use. Recently, a poisoning axe has been developed and used successfully in the South but there is still much room for improvement. Recently a series of translocating poisons, of which "2, 4 D" is a well known example, have been developed. While potentially extremely useful in cover control, they are as yet too new to have been thoroughly tested for effect on many trees and shrubs. In New York State the District Wildlife Managers are familiar with test results as available and may well be consulted before cover control measures are decided upon. Grazing Woodland grazing is usually highly detrimental to grouse. \^Tien long continued, it reduces the variety of the plant life present. Grass is encouraged. Undergrowth and the lower branches of the trees are closely trimmed. In a heavily pastured covert, food and shelter for grouse are usually so deficient as to make it impossible for the bird to survive for long. The parklike look of grazed woods and the "grousy nook" so dear to sportsman and bird lovers alike never go together. There are times, however, when grazing, under New York State conditions, may prove beneficial particularly with overgrown lands, \loderate grazing, long continued, tends to reduce hardwoods in favor of conifers and may result in producing patches of pure soft- wood cover, as well as in bringing in a few desirable food plants such as clover and apples. Some plants, not palatable to cattle but of considerable importance to grouse, as thorn- IN BACK PASTURES, MANY A CROUSY NOOK IS MAINTAINED BY MODERATE GRAZING 642 IMPROVING AND MAINTAINING GROUSE COVERTS apples and blackberries, will often thrive where competing vegetation is removed by grazing. Back pastures, where lightly grazed, have produced many a spot locally famous for the con- centration of i)artridges found there in the early fall. Tangles of thick growing species or of dense underbrush may be opened up by light grazing. Even in some woodlands, restricting or stopping grazing, after allowing it to thin nut the understory, will often help establish new growth beneficial to grouse. As with fire, the secret of successful use of this method lies in knowing when and how to control it so as to bring about the desired result. Where conditions permit its use, moderate grazing is one of the cheapest wavs of retarding succession and of reopening overgrown fields where the vegetation threatens to become too dense to well serve grouse needs. Fire Expertly handled, fire, too, will thin out the vegetation and set back the succession. Other- wise it is likely to cause altogether too much damage, both to soil and to the food and shelter to warrant its use. Nevertheless, it should not be ruled out of consideration as a method of altering existing cover. Where proper controls are possible, light burning represents a reasonably inexpensive and effective way of temporarily eliminating a patch of undesirable species in an overgrown field or of removing undesirable undergro^vth in a woodland. As a method of maintaining small summer feeding units, its possibilities have long been overlooked, principally because of the difficulties of keeping it under control and of the complex public relations involved. Some pertinent effects are discussed shortly: others are noted in Chapter IV. In setting back the succession of plants and trees to an earlier stage, the choice of method is, of course, dependent upon the results desired and the indi\ idual situation to which it is to be applied. Here, again, experience is the best teacher. Producing and Mmntalning Prodlctive Cover The ways of changing existing cover, to produce better conditions for grouse, remain the same over the entire range of the bird. Their application is a problem to be skillfully worked out, for it may differ even within a region. On the basis of experience in New York, a few suggestions for developing and maintaining the types that fulfill one or another of tin- four major grouse cover requirements may jirovc liclitful. Fall Feeding Grounds The cover types that serve primarilv as fall feeding grounds have their origin largely in fields or pastures, once cleared but allowed to seed in with shrub and tree species. Slashings or openings caused by windfalls also serve this purpose to a considerable extent, l^sually. such areas contain a large proportion of wind-sown species such as aspen and birch, and of those whose seeds have been carried into the fields bv birds or mammals. Included in this latter group are cherries, thornapples. dogwoods and \il)urnums together with an occasional sprinkling of apples if the land was once pastured. Pine or spruce, too, here find conditions favorable for germinalioii and will often become establisliod if seed trees are located tn wind- ward. The ideal arrangement of trees and shrubs in a fall feeding ground is a scattering or clumi)- ing of such species interspersed with patches of low herb growth including clover and berries. * S«e page 232. ALTERING EXISTING COVER 643 Cordtner Hump IN THE EARLY STAGES, SMALL PATCHES OF FALL FEEDING GROUNDS MAY BE MAINTAINED RATHER EASILY WHEN ALLOWED TO GROW UP, CONSIDERABLE CUTTING IS NECESSARY TO RELEASE FOOD SPECIES FROM THE COMPETITION OF THEIR TALLER NEIGHBORS. WITHOUT THIS HELP, APPLE AND SUMACH (indicated BY' ARROWS) WILL SOON BE CHOKED OUT r>H IMPROVING AND MAINTAINING GROUSE COVERTS Improvement practices, therefore, consist of periodic release to free the best developed trees and shrul) cliinip-i fiirnishing good fall food from the competition of their more vigorous but less desirable neighbors. Apples and thornap|)Ies, for instance, may be over-topped by trees which normally grow taller. A few strokes of llie axe or poisoning tool may help to keep many of the former in fruiting condition for years. It is also productive to trim u|) old apple trees, relics of the past, but ofttimes capable of })roducing sought after buds and fruit for many years, if given a little care. It is wise to have a backlog of such projects against the time when other activities are slack. Limited grazing is also an aid in maintaining the fairly open character of the types mak- ing up fall feeding grounds. Too much j)asturing, however, tends to make islands of the tree and shrub clumps by maintaining close-cropped grasslands between them. Where fall feeding grounds are in imminent danger of growing into second-growth wood- lands, a more extensive cutting, girdling or poisoning of the larger and less desirable trees may be necessary if the type is to be perpetuated. In fact, of all the types making up grouse cover, none presents a more difficult maintenance problem than do overgrown lands. Nor is the problem made easier by the fact that one must be able to recognize a great variety of valuable shrubs in addition to the usual tree species with which any competent woodsman is familiar. In heavily wooded areas, where fall feeding grounds are not present, the development of clear-cut units has proven valuable, since some species furnishing good fall food will always be found there. The method of developing these is described later in this section. Spring Breeding Grounds Grouse have been known to nest in every type of cover from shrub-dotted fields to the cen- ter of a small patch of dense forest. Most, however, prefer open woodlands where the under- growth is not too thick. The presence of conifers in the immediate vicinity apparently makes little difference. Most birds choose a site within a hundred feet of an opening, such as a woods' edge, a piece of cut-over land, a w^oods road or a windfall, where succulent vegeta- tion will provide food and shelter for the prospective brood. Any second-growth or mature woodland, if it is not too thick, nor composed mainly of conifers, may furnish acceptable spring breeding grounds. Even fall feeding grounds are occasionally used. The characteristics described here and in greater detail in Chapter III* are seldom difficult to find in grouse coverts throughout the Northeast. The common practice of utilizing farm woodlots for light pasturing, for furnishing the winter's supply of wood and for an occa- sional crop of timber for farm use or for sale, has, in general, provided ideal grouse nest- ing cover. In extensive forest lands, action by insects and disease often results in setting up somewhat similar open conditions. Where the forest has not been lumbered or burned over for a long time, the stand may sometimes be so thick as to make it desirable to selectively lumber small areas or to cut deformed trees, here and there, to open uj) the crown cover. The most desirable conditions seem to result when the forest canojiy is not allowed to occupy more than 60 to 80 per cent of the available space overhead. This allows sufficient light to filter through to encourage a not-too-dense undergrowth. ♦ See pafe 126. ALTERING EXISTING COVER 645 Gardiner Bump FORTUNATE IS THK COVERT WIIK II SUPPORTS A SCATTERING OF APPLE TREES ALONG ITS EDGES A BRUSHED-OIT \\ OODS' ROAD IS ATTKACTU E ALIKE TO NESTING BIRDS. BROODS AND ADULTS 646 IMPROVING AND MAINTAINING GROUSE COVERTS Occasionally one will be faced with cut-over lands which have grown up into a dense jungle of large sprouts. In accordance with good forest and game practice, these may well be thinned out especially if other nesting cover adjacent to openings or woods' roads are not available. In carrying out such cultural operations, it is wise to give preference to trees furnishing food and shelter as well as to those worth saving as timber producers. An excellent, though temporary combination of spring breeding grounds and summer feeding units may often be established, in the Northeast, by cutting out an occasional, usually overmature, widespread- ing "wolf" tree. This encourages the light-loving herbs and berries to take over the forest floor which, otherwise, is too often bare. Where den-loving wildlife, such as squirrels and raccoon, are desired, hollow trees may be girdled and left in place. Woods roads also help to break up the forest c()^pr and provide small openings along which partridges prefer to nest. W inter Shelter Unless a woodland is heavily grazed or recently burned, adequate shelter for grouse is usually present except in winter. At that time clumps of conifers or, in their absence, thick groups of evergreen shrubs such as mountain laurel, mav provide congregating places for the WUKKK WINTKR SIIF.LTKH !,>; .SCARCE. A .SPOT SUCH AS THIS MAV WVAA. UK THINAKI) TO KNCOl K- AGF THK K.VERORKENS BEN£ATH iiirds. W IiIkhiI llicse, at lea.xt in tin- Northeast, grouse are apt lo experience serious difli- culty in a\ol(iing predators while ki'c|)inp comfortable. WIicti ((irulitidns allow, lliev will resort to snow roosting as a substitut<' but here, also, lhe\ arc liable lo attack. Hardwood ALTERING EXISTING COVER 647 covers and fall feeding grounds, bereft of such evergreen shelter, unless unusually dense, seldom attract many birds in New York while snow covers the ground. Where conifers occur naturally in clumps or blocks scattered through the woodland, little need be done except to maintain them and encourage their reproduction. The occasional opening up of the forest canopy will allow young evergreens more light. Where conifers are not present, a fairly acceptable substitute may result from establishing a scattered series of clear-cut units, each occupying from one to two acres. The tangles of sprout hardwoods, vines and berries which grow up following cutting may be depended upon to provide some shelter until the crown cover closes. The interplanting of conifers to provide winter shelter is discussed in the section on Conifer Plantings. Summer Feeding Grounds No other summer feed approaches the omnipresent raspberries and blackberries in popu- larity. This may help to explain why so many grouse, old and young, spend much of July, August and September in the cut-over lands and along the feeding grounds that stretch out from the woods' edges. Windfalls and selective lumbering, also, often let in sufficient light to encourage small islands of these berries amid the com|)arative bareness of the surrounding floor. Thus is produced a typical summer feeding ground jjattern rich in herbs, shrubs, leaves and berries that produce food and shelter in endless profusion. The genesis and development of the overgrown field or woods' edge type has just been discussed. Of the three major types producing summer cover, this, while always popular, is, however, least sought out. More attractive are the recentlv spot-lumbered woodlands, the result of cutting individual trees, or clumps, here and there. \\^ind. disease or insects, by attacking now and then a spot or a species, tlirougliout the forest oflcii pio(hicf much the same result. At the top of the list, however, at tliis season stand the small cut-over areas or slashings. This is natural for if one studies such situations he will find there the greatest variety of food-bearing species and the most birds. The name "cut-over" indicates the way in which such summer feeding grounds usually originate. The farmer, intent upon securing wood for farm use. and the lumberman, by removing small blocks of salable conifers and hardwoods, may each be doing the grouse hunter a real service. Again, the same result may originate through natural agencies, such as fire, insects or disease. Unless controlled, however, the area affected is usually far larger than necessary and is seldom properly situated. It is probable that in extensive woodlands a few sunnner feeding ground units will need to be opened up. Suggestions concerning their location, size, number and rotation have been discussed in the previous chapter. Here we are concerned with the actual process of estab- lishment. There are at least three ways of accomplishing this. One may establish what is colloquially known as a jungle slashing bv cutting the trees and letting them lie where they fall or by lopping off only the larger limbs from the trunk. Or the undesirable trees may be girdled or poisoned and left standing. Either practice is apt to create a fire hazard over a small area and should be used onlv where the cost, as indicated in table 96. must be kept as low as possible and the potential logs or cordwood are not worth removing. Though the appearance 648 nil'IWVING AND MAINTAIMXG GROUSE COVERTS of a small iiiiil. llui.< treated, is not |)re])()sses?iiij;. lijiht-lox ing: vegetation s i takes over the site and the grouse find little diflicult) in making; full use of it. Where inexpensive labor is available or wood is salable, the trees iiiav be cut. the logs sold and the brush piled (ui wideh -spread large limbs laid on the ground. Such brush piles SLASH LANES, FROM 30 TO 50 FEET WIDE PROVIDE EXCELLENT SUMMER FEEniNC r.ROlNn? for broods and fall FKEDINC. \REAS for \DILTS fuiiiish sdiiir immediate cover which. allliouj;h grouse arc mil particiilai l\ partial to the shelter thus pro\ idcd. other wildlife will use. By piling and burning the surplus brush, one may encourage the seeding in of desirable species such as |)in chcrrv and aspen that f(dlow fire. IXHI.K '((.. \l'l'li()\l\l\ll-; COST OF E.S'r.\HM>IIIN(, VM) \l\l\rVIMN(i CLi;.\n-ciiT DiNirs by naiuoi s mi:iii()I>s L.NDER NEW YORK. CONDITIONS Typi' iiiitl tiifrlinil lit r.shihlish Cost prr nrrr iif piTitiilic iiiiiinl(Miiiiii-i* JiiriKl<* slushing — lurf^r timhs li»|»|ifi| $l5-$25 :.- 10 60- an S In lie left. fa\or species producing mast, cover, palatable buds and leaves. Do not encs llic advantage of low initial maintenance cost. On the other liand. inatn broad-leaved species sprout prolifically. \\ here the trees to be removed arc mostly hardwoods, girdling or poison- AN IDEAL SMALL SLASHING. CLEAN CUT. IRREGULAR IN OUTLINE AND WITH EVERGREENS NEARBY 650 IMPROVING AND MAINTAINING GROUSE COVERTS iiig may substantially ictanl cidwii cIumiic Ii\ inliiliiliiip suckering. British gamekeepers have long used coiilrollud Imrniiig as a means u{ assuring a fresh growth (pf heather on grouse; moors. \^ ith adequate safeguards, the same principle may be applied to keep open our own small, cut-over units in nmeh the same way as a railroad bums the brush from its right of way or a farmer may l)urn over a field. The fire must, however, be kept on the surface lest the soil be harmed or too many desirable food species be killed and grass encouraged to take their place. The exceedingly dry periods, when fire is likely to get out of control, as well as the months of May and June, when grouse are nesting or the broods are small, obviously should be avoided. Where grass is likelv to follow repeated removal of the forest cover, a rotation of units to be used as summer feeding grounds is called for. Other reasons for this practice have been discussed in the preceding chapter. Correlation of Grouse Cover Improvement Work vi'ith Other Forest Practices In carrying out management, some conflicts in desirable forest and wildlife practices are unavoidable, but the concessions, which must be made to encourage the production of several crops concurrently, are seldom as serious as is generally supposed. In developing the covert, full advantage should be taken of forest management operations such as thinning, improvement cuttings and the harvesting of wood products. In terms of dollars and cents intensive cover development for grouse alone is apt to be a costly under- taking. Combined with other forest improvement work, it can often be made to pay its share of the total expense. Briefly, the steps by which this may be accomplished are the following: 1. Determine what otlier products can be produced. 2. Find out for which of these a profitable market exists. 3. With the above in mind, select the other crops \sith due rejiard to their efTect on grouse. 4. Enlist technical assistance, where available, in determining how best to encourage and when to harvest each crop. 5. Carry out forest stand and game cover improvement practices when labor costs are low. Since grouse coverts are largely wooded, one thinks naturally of lumber, pulpwood, acid wood, ties and firewood, as the possible subsidiary crops. The local market demand for these is not always easy to determine. Softwoods of siifTicient size to produce hmdier are usually salable. The demand for hardwoods is less easil) i)redietal)le. With the de\elopment of new processes, an increasing number of pulp mills are purchasing both hardwoods and softwoods cut in standard eordwood lengths and with a diameter limit as low as four inches. Some mills accept only peeled bolts, others can use the wood with the bark left on. In New York State the market for acid wood is limited largely to the western Catskills. Ties are always in demand though contact must usually be made with the district purchasing agent of the railroads in the vicinitv to find out just what is salable. Likewise there exists in most communities a market for fuelwood, although in limited quantities. Naturally the periodic harvesting of ain of these crops will alter the habitat for grouse to some extent. On a long-time basis lumber iiroduction is quite eonipatilile with the mainte- nance of high grouse populations providing the basic requirements of the bird are met. The ALTERING EXISTING COVER 651 following arc a few general suggestions whicli may help in this respect. 1. Where practical, woodlands should be managed on a sustained yield basis. This implies that the cutting is to be spread over a number of years, preferably by a periodic lum- bering off of crop trees. 2. The areas to be cut in any one operation should be kept as small as practicable. 3. Avoid encouraging stands composed of but one or two species. 4. Where possible, maintain mixed stands composed of hardwoods and conifers rather than a pure stand of either. 5. Leave sufficient conifers, preferably in clumps, well scattered throughout the woodlands, to furnish escape cover and winter shelter for the birds. 6. Encourage uneven-, rather than even-aged, stands. 7. Follow the practice of periodic selective cutting of crop trees rather than clear-cutting large areas. 8. Always keep the basic cover requirements of grouse in mind. Carry out the timber harvest so as to increase rather than decrease the number of good grouse habitats. The management of a covert to produce pulpwood as the main timber crop is likely to ha\e serious repercussions on the grouse population unless the cut-over areas are fairly small and are well scattered. This is seldom considered to be practical. On the other hand, where a market is present for pulp, there exists a golden opportunity to make thinnings, cut in connection with the maintenance of overgrown lands and woodlands, return a profit. On New York's game lands several thousand acres of woodlands have been thinned in accor- A COMBINATION SUCH AS THIS OFFERS A CHALLENGING OPPORTUNITY TO PRODUCE LARGE CROPS OF BOTH GROUSE AND TIMBER dance with good forest and game management practices, the wood thus cut being transported up to 80 miles to pulp mills for sale at a price sufficient to cover the cost of the entire operation. Much the same situation exists as regards acid wood as a crop to be cut from grouse cov- (.52 nirnoi i\(; i\f) 1/ //v7//\/vc; cRorsF. coverts erts. \\ here lulliiijr can lie Wdrknl iriln llic i;iiiri;il plan fur (lixclupmi'ril and niairileiiance of grouse cover ull is well — but this is seldom the case. Perhaps the most satisfactory situation occurs when it is possible to combine fuelwood and grouse as joint crops. Here it is seldom necessary to cut large areas at one time and the operations may usually be so scattered as to make them fit excellenth iritii the game cover iin|)rovement program. It is seldom necessary to make the final dciision. either as regards the croji to he encour- aged or the ( ultiiral practices to be carried out. alone. HEAVILY I'KLiNKI) I'l.WT ATIONS AM) (.KOI SK ARK \(n t.oM )■ \ liHI.i; If a landowner is in need of leclniical assistance in Tiianaging his woodland, there are sev- eral service groups to which he nun tnm. In nian\ stales there is the Extension Service of the Slate Colleges of Agri( nllnrc ant Ian(l(iwnir> to prodni c linilx'r and game crops on a sustained \ ield i.asis. In some counlies tiiere are also Farm Foresters who s])eciali/.e in assist- ing private owners of forest lands. A discussion of coverl problems with an\ of them is likely to ])r(i\(' highlv profitable. Forest Stand Ini i>i(ivcnifiil II oik Every overgrown licld and woodland needs to be thimii-d occasionalh to release the crop trees from undue competition, slinndalr tlii-ir Limwlh and niainlain a doiralile composition and arrangement of the cover. ALTERING EXISTING COVER 653 The word "desirable" here coiiiioles Irces and shrul).s furnishing food and shelter for wild- life as well as those that provide forest products. In England, such thinnings are commonly employed to maintain the open character of the young stand so desirable for game produc- tion. The same practice, applied in New York, can often be made to encourage much sum- mer and fall feed for grouse, and to develop excellent winter shelter and spring breeding grounds as well. How best to carry this out to serve the needs of both forest and game has accordingly been carefully considered. The brief guide, currently in use on the State game management areas, is reproduced on p. 777 to p. 7o2 in the Appendix. The pruning of softwoods is one practice which it is well to use sparingly on grouse areas. Where conifers are abundant, as in a platitation. the lower limbs on possibly 1.50 crop trees per acre can be cut without seriously impairing the shelter value of the stand as a whole. Where this is carried to the point where the stand liironies open beneath the crown cover, one finds grouse notably absent. In carrying out forest stand iniproxcnicnt operations over a large area, it i.s .-cldom neces- sary or desirable to open up the woodland to such an extent that parklike conditions result. Where heavy cuttings are needed to create the jiroper com|)osition. it is suggested that they be spread over a number of years, thus encouraging lielter tree form and lengthening the period in which abundant summer food is available. If llic existing woodland cover is of little value, it may be considered a nurs(> crop ;ni(l In- xi bandied as to cncouraiic llic reproduc- tion of the more valuable species. As the woodland becomes older, good foic-si piaclicc calls for llic rcmliuiicd rcnunal of dead or defective trees, as well as for the fuilhcr release of those thai arc lo provide the final timber crop. Here forest stand improvement practices can be carried out in a manner to encourage wildlife b\ so opening up the forest canopy as to maintain beneath it a fairly abundant undergrowth. On New York game areas this has been aci-oniplishcd b\ cutting enough trees so that the remaining crowns occupy from .50 to 00 per cent of the available space. Even if the stand is of inferior quality, lo open up the forest cover further may encourage erosio?i. stimulate grass beneath, and forfeit the advantage of a canopy sheltering the reproduction so desirable if one is to maintain a sustained timber yield. On the other hand, to leave too many trees is to encourage a relatively bare forest floor acceptable, per- haps, to grouse as a nesting area, but deficient in sununer and fall food as well as in desir- able young growth. As the forest matures, the method of harvesting the timber mav likewise increase or de- crease the productivit) of the area for grouse. The selective removal of crop trees creates small openings thereby maintaining sufficient shelter and food for the birds. Relatively clear cutting even to a diameter limit of six to eight inches, over large areas, on the other hand, is usually detrimental by creating a super-abundance of summer feeding area. To prevent this, scattered small slashings are always more desirable, where practical, than is a single large lumbering operation. By giving some thought to the maintenance, on each acre, of half a dozen trees providing food and shelter and by encouraging their reseeding and growth, the game-producing attributes of the stand may be maintained. By skillful cutting, it is often possible, over a period of years, to secure an even better distribution of species which collec- tivelv will produce a forest and encourage a grouse crop. Other ways of increasing the crop of wildlife and of forest products will suggest them- >^- 654 IMPR(WING AND MAINTAIM\G GROUSE COVERTS selves to the skillful forest manager interested in oi)taining a maximum harvest from his lands. Those mentioned here have been found practical in New York. It is recognized, of course, that in other regions it will l>e necessary to adapt such practices as lit local conditions and environments. Hut if one is sufficiently interested, there is little doubt that it can be done. Control of Forest Insects and Diseases The measures currently in use in New York for controlling forest insects and diseases do not seem to affect grouse much one way or the other. True, weeviled white pines are apt to provide better shelter for birds than do straight, fast-growing trees but the difference is seldom sufficiently pronounced so as to make it inadvisable to carry out control measures on young trees. Terminal shoots, infected by the white pine weevil, may usually be recognized by the wilting of the terminal leaders. Control consists of cutting the shoot below the hole during the month of July and burning them. The eradication of gooseberries and currants, either wild or cultivated, from within the covert, to prevent the spread of the white pine blister rust is important in those areas where this destructive disease is prevalent. Much concern has recently been expressed over the possible harmful effects on bird life of spraying woodlands with a weak solution of DDT to control insects such as the gipsy moth, the spruce bud worm and the pine saw-fly. While young grouse feed to a considerable extent, on insects during the first ten days following hatching, where concentrations of one pound per acre have been sprayed from an airplane, no harm to grouse, old or young, has been noted. The indications are, however, that larger applications may prove dangerous. DEVELOPING FOOD AND SHELTER BY PLANTING* In many cases, the habitat may be improved by planting additional shelter and food pro- ducing cover types. How this can be done will be determined by the existing conditions on the area. The methods to be used on open fields will differ from those needed for cover already established, while varying deficiencies within that cover will call for particular plant- ing techniques to improve them. Basic Principles Controlling Planting In general, at least portions of the improvement must be accomplished by planting seeds or seedlings. There are certain basic principles that should be followed. They are: 1. Determine the proper establishment method required. 2. Test for the most important soil factors, such as its f(Tti!it\ l<'\cl ami jnoislurc char- acter. Site requirements ior trees and shrubs, recommended for planting in New ^ork. are given iti table 97. 3. Select species suitable to the climate, light and shade, and soil conditions. These are shown in table 97. 4. Secure suitable seed and planting stock. .S. Care properly for planting stock. 6. Use the proper planting methods. • By Frank C. (Idminttcr, DEVELOPING FOOD AND SHELTER BY PLANTING 655 Determining Cover Eslahlislinirnl Methods to he I srd The methods by which the cover types shoultl lie estabHslied arc the (usl coiisiileratioii. The character of the soil is the major limiting factor. The simplest and cheapest way, pro- viding it works out satisfactorily is to let nature take her own course through the orderly process of plant succession, while the most difficult and expensive one is to set out each woody plant. In some instances, there are intermediate stages that may be employed whereby one can avoid complete planting of the area by assisting the establishment of desired types by cultural operations that promote the growth of certain species and groups of plants, while retarding unproductive ones. TABLii 97. USES OF TREES, SHRUBS AND VINES FOR PLANTATIONS. AND THEIR SITE REQUIREMENTS Outstanding uses of the plant or its product Moisture tolerance Fertilily reqiim*- menU Shade tolerance Species* Q 91 ■5 J 2 s .1 Ij s 1 Conifers: Red Pine White pine European larcli A Norway spruce Winter shelter, mlllwork. Xmas trees, t-onstruction lumber, boxes, pulp. Winter shelter, construction and millwork lumber. boxes, patterns. Xmas trees. Construction and millwtirk lumber, boxes, posts. Winter shelter, pulp. Xmas trws. construction lumber, interior trim. Winter shelter, pulp. Xmas trees, construction lumber, interior trim. Winter shelter, pulp, Xmas trees, construction lumber, int(*rior trim, also musical instruments. Winter shelter, posts, construction lumber, bouts, cooperage. Kood at all seasons, furniture lumber, fuel. Winter and spring fo(Kl, fuel. Maple syrup, furniture lumber, fuel, shoe heels. flooring. Fall, winter and spring food. fuel, farm im piemen t.s. Spring and fall food, construction lumber, ties. flooring. Spring and fall food, construction lumber, ties, flooring. Spring and fall food, fuel, flooring, construction lumber. Winter and spring food, oil, fuel, woodenware novelties. Winter and spring foi(iii. (Jiii' iiiii>l cxiiininr ihc i'\i>liiii; cuiHlilidiw and dclcnnine what llic |)riilpal)ililit's arc for success b\ tilt' \aiiciU5^ tiiethods. \aluially. tlmsc selected will \ar\ according to tlie t\ |)e of habitat desired on am given location. Ihus. if the design calls for fall feeding grtninds. one must consider the |)ossihilit\ of olitaining the desirable shrub cover. If seedlings of the required species will establish themselves by natural secl partial planlini;> >liiiiilil be made. Where the habitat alreadx exists, the nielhod> to be used will depend upon what changes and improvements are needed to produce the required habitat. Some of these methods have DEVELOPING FOOD AND SHELTER BY PLANTING 657 already been discussed under the section on altering the existing cover, wherein the making of openings and clearings is described. These may be used for interplantings and estab- lishing herbaceous cover types. Securing and Caring for Plaiiliiig Slocli Planting stock for extensive revegetation work must, necessarily, be rather low in price in order to make the operation economically feasible. At the present time, state and federal nurseries produce a good many of the species recommended for the different types of plant- ings and make them available to the public, in various ways, at a reasonable cost. The i)ro- duction of state nurseries is predoniinanth of coniferous species, thus making available the material for the greatest portion of ])lantation work. It is often possible to obtain suitable planting stock from pri\ate nurseries. Material known as "lining-out stock" in commercial inirscries is often purchasable at a reasonable cost. Gen- erally speaking, a desirable hardwood seedling of one or two years nurser\ growth is from six to fifteen inches high and uilh a root s\stem nearly as large. Care in selection of conifer stock for use in interior plantings is very important. The stock should be large and thrifl\. For the species recommended, a four-year old nursery grown plant is usually best. Foresters call them "2-2 |)lants" to indicate that they have been grown in a seed bed two years and in a traii>|ilaiit bed an additional two years. Planting stock should be carried in a pail parlh filled with water so that the roots are not permitted to dry out. Portions of il that can not he ])lacc(l in the ground innncdiately upon removal from the nurscr>. should In' liccicd-in in a damp, shady, well-drained spot. To do this, a trench is dug deep enough to accommodate the root s\stems. about a foot wide and long enough to take care of the tnalerial at hand. The bundles of seedlings are loos- ened and laid with the roots in the trench and covered firmly with soil. The plants are kept moist by wetting and covering with burlap. 'Hicn shduld lie removed from the heel-in beil only as needed for immediate planting. Some wildlife managers may wish to inopagale their own seedlings. This is an immense subject by itself which eamiol be covered here. There are numerous texts on the subject and a wealth of experience, in both public and jirivate nurseries, is available. Most of the information from these sources, at least in so far as it concerns reforestation use, is con- fined Id the conifers and a \ery few hardwood trees. For those who desire to propagate |)lants for grouse habitat. particularl\ the shrubs and hardwood trees, the specific data needed is summarized in table 186*. Most of this material was recorded through the courtes\ of th(> Soil Conservation Service and with the assistance of Robert B. Thornton, formerly Cliief of the Nursery Division of the Northeast Region. I'hiiiling Methods In placing seedlings in the ground, considerable care should be taken in order to assure good sui-vival and growth. In most soils, the best planting method is that known as the center hole technique. Uy using a mattock or other similar tool, at least a square foot of the sod cover is removed and a hole dug in the middle of this "scalp" deep enough and wide enough to accommodate the seedling without cramping its roots. The seedling is held in the center of the hole with the root collar at the ground line level and the soil is then placed firndy around the root and tamjied down at the to|). usually with the foot. * Set' Apppntlix. p. 883. 658 JAJFKOyiNG AND MAINTAINING GROUSE COVERTS Direct seeding of nut species is arranged as though the seedlings were being planted. Sod scalps are first made. Then twn nuts are buried in the center of the spot about twice as deep as the diameter of the seed. Rodents cause considerable loss to plantations of acorns and nuts. For this reason, plantings should be kept away from the edges of existing woodlands in so far as possible. Greater success will be attained when these plantings are made in years when squirrels and mice are not excessively abundant. No satisfactory material has yet been developed to repel rodents from either nuts or seedlings. PLANTING SPRUCE BY THE CENTER HOLE TECHNIQUE Seeds may be planted dircctlv into newly ])lowed furrows. As a precaution against unnec- essary soil loss, the furrows should be plowed on level lines, that is. crosswise to the slope. For large shrub niul tree species, tiic furrows should be about six feet a])art and for small shrubs, about four feet. For herbaceous cover, the seeding may be done by sini])]\ disking with the usual farm disk- ing-harrow in order to prepare a good seed bed. The germination of conifer seed is stimulated by exposing the Miiiici;il soil liy ploughing the furrows. When seed trees are present, natural seeding in the furrows will usually suffice but, if not. then seeds of the desired species may be planted in iheiii. This practice is still in the experimental stage and will require a great deal more study before the teehnieal details iiMjiiiri'd to obtain successful growth cmii lie lui'diilid willi ;in\ a<'ciira(\. The jxtssibilities of establishing hardwoods liv similar direct seeding methods are even less well known today. Certain types of liariiwond shrubs can be grown by direct seeding on DEVELOPING FOOD AND SHELTER BY PLANTING 659 seed beds prepared in a manner similar to that for herbaceous cover. None of the shrubs that are of particular use to the grouse are among those that can be established with assur- ance. Conifer Plantings Most of the conifers are adapted to growing in soils that are acid and low in fertility. Like most plants, however, they will do better in more fertile soils. Therefore, since the hardwood trees and shrubs are more demanding in this regard, it is a good plan to depend n':^- ^'^■, k-»?^-'' - .^^"^ ' MOST HARr)WOOD TREES AND SHRUBS RESPOND BEST WHEN PLANTED DIRECTLY INTO NEWLY PLOWED FURROWS upon the c-onifers for revegetating ihe poorer sites. One nmst not expect the most vigorous growth under these conditions. The species of conifers best suited for winter shelter are hemlock, white pine, red spruce, white spruce, black spruce, Norway spruce, red })ine, and northern white cedar. It is unfor- tunate that hemlock is difficult to produce in nurseries for reforestation use and is not de- ])endable in field plantations. Under most circumstances, the wildlife manager will depend upon the red. white and Norway spruce, white and red pines and northern white cedar, for the greater portion of his winter cover. Another desirable conifer in the mixture is European larch. While not an evergreen, this tree provides a necessary element in the winter shelter composition bv developing a useful herbaceous ground cover, often lacking with many other conifers. If other species of pines, spruces or firs are available and \ aluable for their wood products, they may be included in the mixture. In all plantings for winter shelter, it is advantageous to 660 IMPROVING AND MAINTAINING GROUSE COVERTS use a mixture of several species rather than jjiantiiif; in blocks of a sinfrle vaiict\. The species arran{;cnient tiia) lie either as a random selection from ])ail mixtures or in continuous rows of each kind. Where the development of winter shelter, in hardwood stands dehcient in conifers, is re- quired, it is often necessary to interplant. Within the woodland, this should be done in ex- isting open glades wherever possible, or openings should be made to pro\ ide the necessary sunlight for adequate growth. This may be accomplished when har\esting the wood or by I'. S. Stiil Ciinarriittion Service W HITE PINE. INTERPLANTED IN AN OPENING IN SECOND-GROWTH HARDWOODS, HERE I'liOVMlES Ml CM NEEDED WINTER SHELTER FOR GROUSE silvicultural thinnings, as |)reviouslv discussed.* In cover thus opened, there still may be considerable shade and. therefore, sliadc-lolcrant species, such as white or red spruce, are recommetided. If the crown canopy is sufficiently open, so that the site is not more than half shaded, then whhc ])inc. Norwax sprmc and northern white cedar may also be used successfulh . ('uttings of hardwoods on sites intended for inter|)lanlings will gcncralK result in con- siderable spiouls and stinnps. These vigorous shoots are likcK to result in excessive com- petition and shading for the newly-planted seedlings. I'nder these conditions, it is suggested that the slumps of the cut trees be poisoned"' in order to insure adequate sunlight for the ra|>id eslablishmcnl of ihe evergreen trees. Spacing for inlerplantings must be judged on the conditions at cadi Idcatiiui. Ordinarily, the usual six to eight foot plantation spacing may be used. Where the openings are small and scattered, it ma\ be desirable lo set the platits somewhat closer together. IJVUDUOOD TuKi: I'l.VNTINCS W lien planting i^ needed for the de\elopin<'nl of s|)ring nesting grounds, one must use • flisrilimion of AllrriiiB |I»* Kxtittinc Ciivrr. |t. iiii>ning, p. (110. DEVELOPING FOOD AND SHELTER BY PLANTING 661 considerable care in the selection of the sites. Hardwood trees usually require a more fer- tile soil for successful establishment than do the conifers. It is fortunate, however, that, in most portions of the Northeast, natural plant succession will develop some hardwood tree species. In most cases it is only necessary, therefore, to plant a portion of the area to attain a better balanced hardwood stand. This provides an opportunity to select the best ground for setting out the trees where artificial planting is needed. WHERE THE CROWN COVER IS KAIRl.Y OPEN A.N UNDERPLANTI.NC OF SHADE-TOLERANT SPRUCE MAY ALSO FURNISH WINTER SHELTER A mixture of species is just as necessary in hardwood |)lantings as with the conifers. Those listed in table 186* have been selected because of their particular value to spring nesting grounds. Others, such as the walnuts, hickories and chestnuts, while not of particular im- portance to grouse as a source of food, are of sigiiilicance to a number of wildlife species. These large seeded trees, as well as the oaks, may be established by planting the nuts instead of the seedling trees. There are a number of other species of hardwood trees that are suitable to plant but which are not of special use to the grouse. However, one may wish to add some of these to the mixture for their value as wood products. Among these are tulip poplar, white ash and basswood. Shrub Border Plantings As previousK mentioned, summer and fall feeding grounds usual!) are best combined in a single strip. In most instances, such a stri|i is a border of the cover. A minimum width (if about 25 feet is needed in order to maintain this plant t>pe. The width may be increased up to approximately 100 feet or until it constitutes ID to 15 per cent of the habitat area where this cover is particularly needed. * See Appendix, p. 883. 662 IMPROVING AND MAINTAINING GROUSE COVERTS Plantings of the large shrubs and small fruit-bearing trees should be made next to the wood- land trees. The spacing between the seedlings for this portion of the border is recommended to be 6 feet. The small and medium sized bushes may be placed in the outer portion of the border and next to the open land. Here the spacing may well be cut to 4 to 5 feet between rows and plants. By utilizing the lower growing shrubs on the outside and the higher ones next to the trees, a sloping border will develop which provides the greatest protection for the woodland and the best conditions for fruit production. The economical establishment of the summer and fall feeding grounds demands that the wildlife manager, wherever possible, should take full advantage of the opportunities for as- sisting nature as a substitute for direct planting. A portion of the strip may be lightly burned over, in scattered spots, with the expectation that desirable species for summer feeding grounds will establish themselves naturalh. Blueberries, raspberries and blackberries, pin cherry and popple all respond well to this treatment and furnish valuable summer and fall foods. Strict control must be maintained since indiscriminate use of fire is very destructive. For the rest of the border, partial planting is often necessary in order to permit the shrubs to become fully established, or dominant, before tree species can get a start. The proportion to be planted will depend upon the need for additional shrubs, both as to the kind and num- ber of them. Since the purpose of the shrub border is to furnish summer and fall feeding grounds, the types chosen for j)lanting should be those which will furnish plentv of required food for these seasons. A majority of the shrubs and small trees that will do this are also suitable for planting. They are listed in table 97 together with the most essential site characteristics of each. Arranged according to their type of growth, they are: Loic shrubs and vines Medium high shrubs Tall shrubs and small trees Bittersweet Panicled dogwood Thornapple Virginia creeper Hazelnut Apple and crabapple Wild rose Scrub oak Staghorn and smooth Fox grape Arrowwood sumach Riverbank grape Highbush cranberry Nannyberry Dwarf sumach Multiflora rose Some of these species are not among the most important grouse foods. Bittersweet. Vir- ginia creeper and arrowwood, for example, do not rank high as grouse food in New York. However, they arc included because thev do furnish a desirable part of the composition of summer and fall feeding grounds. It is a good policy to plant at least four sjiecies in each shrub strip. One or two kinds from each of the three height groups may be selected ami the species arranged in rows so that there will be four or more rows in the entire border. As the sununcr and fall feeding ground develops, a number of tree species will nsualh seed in naturally among the shrubs. If these are permitted to grow, they will eventually dis- place the shrub border and convert it into a spring nesting grounds type of cover. To prevent this, these trees should be eliminated periodically by ae forgotten that occasional light grazing may be beneficial. If it were not for the cash outlay required for woodland fencing it would not be diffirull to persuade most farmers to do it. However, they often are not certain that tlie cost «ill lie offset by the advantages gained. When one considers the improvement to the livestock, the increase in wood products and the benefits to grouse and other wildlife, the ecoiioinie balance is definitely in favor of protection. Control of Dkkr Browsing In most cases, the over-browsing of the woodland understor) is the result of grazing by domestic livestock. However, in some areas, the while-lailetl deer has broufiht about substantially the same results. The solution to this problem is entirely different tlian that created by domestic livestock. Provision of adequate open shooting seasons and bag limits, coupled with enough hunters, to reduce the density of the deer population below the carry- ing capacity of the range is all that is required. The State should have adequate discretion- ary power to insure enough hunting of either sex as may be required in order to ])re\ent the destruction of valuable woodland cover. CHAPTER XVII THE MAINTENANCE OF A GROUSE CROP By Gardiner Bump, Robert W. Darrow and Frank C. Edminster CONTROL OF ASSOCIATED ORGANISMS Control of Predators — Control of Buffers — Control of Disease RESTOCKING DEPLETED COVERTS When to Restock — How to Restock REGULATION OF THE GROUSE HARVEST Measuring the Harvestable Crop — Determining Grouse Populations — The Proportion of the Crop Available for Harvest by Hunting— Regulation of Hunting £i SUMMARY Most grouse coverts will furnish a sustained yield provided a suitable seed stock is maintained. The number which may be harvested safely varies from year to year according to the size of the fall surplus. ( p. 668). While no serious disease condition in grouse has been encountered by the Investigation, har- vesting the surplus population each fall by hunting is a practical means of guarding against it. ( p. 671 I . Selective predator control may be useful on small areas managed for intensive hunting, mainly in reducing nesting losses, (p. 6f'8). The predators of primary concern to the grouse manager in New York are the fox, weasel and great horned owl. ( p. 669 ) . Restocking of grouse will be justified only where habitat conditions are suitable and for one reason or another an adequate seed stock is not present, (p. 672). A rough, but useful, method of estimating the fall population in local coverts is to count the number of broods in late August and correlate this with the average number of birds in each. (p. 676). Knowing the fall population density and something of the comparative quality of an area one can judge approximately the proportion which can be safely harvested, (p. 677). Refuges as a means of modifying the take are recommended only when hunting pressure is excessively high. (p. 679). r.f..", Tin: \IAl\TE\ANCE OF A GROUSE CROP Imporluiil as is tlii' natmc of lli<' co\i'r and its aiTangeiiR'iit. cmmi llic nmsl carefully de- veloped habitat is no {luaraiitcp of sustained grouse i)ro(liiition. Inavoidahly subject to Nature's vagaries, the species will continue to have its u|)s and downs. The number which may be harvested safely will, therefore, vary from year to year according to the size of the fall surplus. Nevertheless, most grouse coverts will furnish shooting during most vears pro- vided a suitable seed stock is maintained. To insure an adequate number of breeders, as well as to mitigate losses in general, one must consider a variety of factors in addition to the design of the coverts themselves. Restric- tion of hunting bv man has long been a basic practice. Oiilv recentlv, however, have we come to realize that, at times, an increase in hunting may be equally valuable. Control of preda- tors and disease, as well as restocking, when employed properly, may' also be helpful to the game manager. CONTROL OF ASSOCIATED ORGANISMS* The activities of associated organisms, from horned owl to bacillus, are a constant influence affecting grouse abundance either directly or indirectly. Suitable control of such forms may therefore be a useful management tool. Control of Predators Whether or not to undertake the control of j)redators on a grouse production unit nuisl be decided for each area individually. The purpose for which it is being managed must be con- sidered, as well as its size and the nature of the wildlife populations present. As has been pointed out in preceding chapters'^, the reduction of predators is by no means the simple solution to the ])roblem of creating grouse abundance that it is sometimes pic- tured. Moreover, it will not be effective if the (pialily of the habitat is not sufficient to sup- jiort additional birds. And one should remember that, in the Northeast at least, a population density among adult grouse of about one to four acres is all that tlie social tolerance of the birds themselves will permit even in optinnun coverts. Furthermore, the cost of such a program will usually limit its feasibility to comparatively small areas or club properties. On extensive tracts, therefore, management funds would generally be spent to much better advantage in habitat improvement. The same is true with regard to refuges, sanctuaries and other units where little or no hunting is done although indi\ idual predators may occasionally become unduly destructive and need to be eliminated. On the other hand, jjredator control may at times be emplo)ed to ad\antage on units handled primarily for the purpose of hunting. Unusual numbers of certain predatory species sometimes occur and need to be reduced. Again, winter influxes of such species as the great horned owl and goshawk may need to be dissipated. But. in the main, control measures will be found useful chiefly in lowering nesting losses as a means of increasing the annual increment which, in turn, determines the harvestable surplus. At the same time, it should be emphasized that, in the final analysis the quality of the habitat is the cornerstone of game abundance and that neither predator control nor restocking nor similar [)raetice9 will take its place. Even where predator control seems a wise procedure, coniplete elimination of such species over anv appreciable area should be avoided. Their relaliini^bip to gnuise is but a small part of their function in the animal conuTumitv as a whole. • By Ruben W. Danow A See Chapter VM, p. 3S0. ami Chaplir IX. |<. 3 also a necessar) jirerequisite. When grouse are fairly abundant, the normal dispersion of the birds from well to poorK stocked coverts is such thai few favor.d)le ones close by will remain seriously underpopulated * Hy Cardinei Bump A S» Chiplcr XI. p. SO-I. REGULATION OF THE GROUSE HARVEST 673 for long. Naturally, as the distance between coverts increases, the likelihood of restocking them by dispersion decreases. While little indicative information is available, it seems doubt- ful that much natural transfer of population is likely to occur when the distance exceeds four or five miles. Such natural movement may obviate the expenditure of effort and money involved in a restocking program, fndividual circumstances may occasionally arise, however, which will justify this expenditure. How TO Restock Providing the conditions outlined at the beginning of this section are met. and one takes due care to avoid the causes of the many previous failures, successful restocking should not be too difficult an undertaking. The birds should be liberated in the best habitat possible in groups of not less than 20 to 25. It is important to repeat the liberation annually for at least three to four years to permit the birds to receive additions to their numbers to make up for a normal loss incident to their establishment. The best period for liberation is believed to be in the summer or early fall, which corre- sponds to the time when artificially propagated birds seven to nine weeks of age could be secured. Wild birds are more easily trapped in the wintertime and are. therefore, usually liberated as adulls. Since they lose weight consistently between early January and mid- March it would seem wise to liberate them either as early in the winter as possible or in April. Unless they are in poor condition at the time appointed for liberation they should not be kept in captivity any longer than is absolutely necessary lest they lose additional weight. Young hand-raised birds may be fed at the point of liberation though their wild-trapped brethren will pav but little attention to such efforts. The carriers from which they are liber- ated should be placed so as to make it diffit ult for the birds to fly upon emergence. This will encourage the birds to remain as long as possibh- in the immediate vicinity of the liberation point. REGULATION' OF THE GROUSE HARVEST* Regulation of the harvest itself in acronlaiue with the relati\e abundance of the species from year to year represents the concluding phase of a management program. In America the taking of non-migratory game is governed by what('\ cr limitations the various state legis- latures see fit to lay down in their game laws each )ear. Within the limits thus established, however, it may become desirable for the administrator as well as the individual game inan- ager to make additional adjustments to fit particular situations. Thus, if the grouse crop should fail on a managed area, it would be logical to set up on it further restrictions than pro- vided by the current state-wide laws. The regulations adopted should be directed toward maintaining a sustained yield. Neither too many nor too few of the species should be harvested. This means that in years of plenty the crop should be adequately utilized, whereas, in years of scarcity, the grouse should re- ceive sufficient protection to provide the greatest possible hunting opportunity and at the same time insure the preservation of a sufficient breeding stock. A reliable means of meas- uring the crop, or shootable surplus, is therefore a jiriniarv requisite to attaining this goal. * Bv Frank C, Edminster 674 THE MAINTENANCE OF A GROUSE CROP Measuring the Harvestable Crop* If the maintenance of a seed stock were the only motive for iiicasurin^' the grouse crop it might be debatable whether the results would be worth the effort. One lould easily "take what the gods send along", shooting birds when they arc aimiulant and passing them by when but few can be found. But the preservation of a seed slock is not the sole reason. Only by knowing their size can above-average crops be utilized to the best ad\antage. And con- versely, such knowledge enables an administrator to make necessary allowances for a vear of poor productivity to prevent an unwarranted open season from cutting into the brood stock. Furthermore, in addition to the Inmtable surplus, a record of po|)ulation trends from year to year is also important in evaluating the results of past regulations. .And, perhaps most significant of all on managed areas, the measurement of the grouse crop is the yardstick by which the productivity of habitat improvements may be appraised. > "**T'iS13lC^:J*'J!"jei^ " '^'ti' ' ' ***** * ' r^ riAtlY & ROUSE ^'1 But it is seldom an easy job to determine, with even fair accuracy, the harvestable crop of any game species. Leopold"' singled out the ruffed grouse as "conspicuously difficult to cen- sus by ordinary methods". The adults do not covey like quail or form packs as do prairie chicken. In the fastnesses of their range, where they are little hunted, they are easily trapped. But where they have come to know hunters their wariness and trickiness is legendary. During the fall and spring, the most significant census periods, many birds frequent heavy cover in which, while often heard, they are seldom seen. Nor are the sex ratios easy to ascertain becau.se the birds look so nnicii alike. While they are not migratory and exliibit no extended seasonal movements, they are mobile over fairly large territories, thus scattering the individuals widely enough to make finding them often difficult. To comiilicate matters further they are predictably unreliable as to their decision to lly at one's approach or to lie close. Furthermore, adjacent coverts of comparable quality, even though exhibiting similar trends in abundance, are usualK at different relative population levels in anv one year. One covert * By Cflrdiner Bump REGULATION OF THE GROUSE HARVEST 675 may be teeming with birds, another at or near a low point. All these characteristics com- bine to make a census notably difficult. Yet the picture is not all dark, otherwise the Investigation could have learned little of the characteristics of grouse populations. The birds are sedentary' in that the adults seem to establish certain territories. When caught near the edge of these they tend to turn back. The fact that individuals or groups of birds are usually scattered makes it easier to avoid duplication once the extent of their territories has been learned. Furthermore the males establish drumming logs which they use regularly in the spring. Likewise, the females nest for the most part in predictable portions of their habitat though their numbers cannot be counted at this time by such an index as the docker droppings (enlarged droppings from nesting birds) of the red grouse of Scotland which commonly leaves them at the same place each day along watercourses. Determining Grouse Populations Reports of interested observers, game protectors and other field workers have commonly formed the basis of abundance estimates preceding the fall hunting season. If ruffed grouse were as predictable in their increase as Scotch grouse we, like the British, might rather estimate the fall crop from the number of breeding birds in the spring. But there are more accurate ways of arriving at such an inventory, although none of the grouse census methods so far developed is without serious drawbacks. The birds are generally too widespread to make practical the expensive direct count prac- tice* used by the Investigation on its study areas. True, such complete censuses are quite workable for small areas providing funds, trained man])owcr and direction are available. On large tracts the time required, as well as the costs, is prohibitive. The next most accurate procedure is to establish a series of sample plots in representative grouse habitats to be measured yearly by the complete census method. This presupposes that the coverts selected have been surveyed in advance and are normally found to carry large numbers of birds. The difficulty here lies in the number of such sample plots which must be covered to give a balanced picture. On a 5,000-acre area, for instance, at least a dozen would probably be necessary. This would make the number prohibitive if a county or a state were involved. Because only highly productive areas are chosen, this method will yield a figure far in excess of the actual population. Used as an indicator of abundance trends, however, it is appar- ently fairly reliable. King"'* developed a somewhat different application of the sample plot method for Minne- sota conditions bv dividing the area to be censused into 40-acre blocks the boundaries of which are followed in making the survey. Figuring the average grouse flushing distance from those recorded each day the proportion of the total area covered is computed by multiplying twice this distance by the length of line covered each time the survey is carried out. The amount of each type of cover over the entire area is established by a type survey. The strips are "walked" at intervals and a record kept of the number of grouse flushed within each type. By comparing the number of birds flushed per unit of each type within the strips to the total of each on the area, an estimate of the total birds to be found in each type may be secured. These added together give an estimate of total area population. * See Appendix, p. 699. (u(y HIE MAI^TEI^ANCE OF \ GROUSE CROP This method is useful where a consideralilc unifnirnitv of cover exists and populations are lelalivcly high, hut otherwise the prohahle error is too great. It is nut suitahle to the well mixed coverts characteristic of the princi])al grouse territory in New York. Its weakness lies in the fact that in this kind of cover such a system of strips seldom intersects enough well ]iopu!ate(l habitats to provide a number of flushes sufficient to avoid large errors in compu- tation. -And to include a larger proportion would require an undue amount of labor since each strip must be worked at least three times to secure a reasonably accurate estimate of the birds occupying it. Relative abundance from year to year but not total numbers can often be estimated after the hunting season by requiring that kill records be kept. An example of these covering 17 years for New York is found in Chapter IX. p. 372. Though many hunters do not report their take accuratelv. the proportion of error remains rclativelv constant thus providing rea- sonably accurate trends. The largest defects here are that such information is not available in time to be useful during the season in question, and that a good kill one year is no guar- antee of the presence of many birds in the coverts the following fall. The length of open seasons, as well as the number of man davs in the field, must also be taken into considera- tion. Other indices of production such as brood and nest counts are occasionally useful where considerable trained labor is available and birds are to be found in sufficient mnnbers. In using the results as indices, however, a correction for the loss occurring between the census and the harvest period is necessary, a difficult job since this figure may vary widely between years. Enough has been said to make it apparent thai no one census method applicable to all con- ditions is to be found. Where a research problem requires an accurate determination of breeding and of early fall po|)nlati(ins and the necessarv expenditure is justified, the complete census based on three closely spaced coverages of an area has proven satisfactory. Probably the most practical method of judging the general ])ros|)ects for the hunting sea- son in local co\erts is through a late August brood count. Enough time nujst be s])ent to cover the area thoroughly, parlicularlv young growth and "edges". Belter results will be olitairicer of broods found, together with tile a\i'rage lunnber of birds in each, will indicate the general population level. One nmst remember, however, that most broods «ill ha\c broken-up ami ihc binls become scattered by the time the hunting season opens. Furthermore, thev will seldom be found in the same jjlaces. \e\erllu'l('ss. if such a practice is contiinied over a period of years in the same area, information secured in this manner should pro\ ide a valuable ind<'x. Over more extensive areas one must be content with estimated abundance. Eor this a combination of a large number of random obser\'ations. backed u|) b\ a brood count in late sunnner or earlv fall on an adequate number of re|)resentati\(' sanq)les of ordinarilv weil- l)o]julated habitats will u>u;dl\ la\ llii' basis for arrixing al an ind<'\ suffiriently good to judge the harvestable crop. The Projiorlion nj llir C.ioj) Iriiihihlf ior fliirrrst hy lluntinf^ If all grouse habilals were fullv stocked, the entire increment each year, save onl\ that recpiired to replace nornud losses, could be removed by hunting. Rut fully stocked coverts REGULATION OF THE GROUSE HARVEST 677 neither occur frequerjtly nor do they usually maintain themselves as such for long, at least in the Northeast. The proportion available for harvest, then, is normally something less than the total increment. Theoretically, as the density approaches the maximum number of birds a covert will hold, an increased proportion of the birds may be harvested. But in actual practice the carrying capacity for many habitats is so low that, even when well stocked (for such coverts), few or no birds may be taken by man without running the risk of reducing the total population to a point where maintenance of the seed stock is endangered. Just where this point lies has not been accurately determined. It is certainly dangerous, however, to reduce the numbers over a sizeable area below one bird to 30 acres of grouse habitat. If it were possible to determine accurately the carrying capacity of individual grouse coverts, as well as the degree to which the fall poj)ulation approached or exceeded that figure, one could then closely estimate the number of grouse available for harvest by the hunter and still leave a reasonable seed stock. Rut the ability of an area to support grouse changes, even from year to year, with the weather, with ])redator-buffer relationships, and with cover succession. Furthermore, the cost of detailed censuses is prohibitive except on relatively small areas. Thus a more practical, if less exact, method is here suggested. Its basis lies in part on 100 liJ UJ75 CO I Q 50 I- Z UJ on a. UJ 100 0 5 10 15 20 25 30 35 40 45 50 55 60 FALL POPULATION DENSITY Cbirds PER lOO acres) u 0 FIGURE 70. APPROXIMATE MAXIMUM ALLOWABLE HARVEST BY HUNTING ACCORDING TO DENSITY OF FALL POPULATION measurements of grouse productivity* made during the Investigation, in part on that inde- finable product of experience called ''grouse sense". In utilizing data from the study areas it has been necessary to make certain assumptions and judgments. The accompanying chart must, therefore, be regarded as the best interpretation the authors have been able to make « See Chapter XU. 678 THE MAINTFA'ANCE OF A GROUSE CROP covering their observations to date. Nevertheless, it is conservative and should serve as a valuable guide to the interested game manager. This sliding scale of allowable take, as shown in figure 70. requires for its application only an estimate of the total fall population in relation to the size of the area involved (birds per 100 acres) and an appraisal of the general quality of the coverts on the area. Oidy data for high and medium quality coverts have been used, however, since it seems unlikely that intensive management of poorer areas for grouse will often be undertaken. Under this yield table, when grouse are abundant or scarce, the take may be correspond- ingly large or small. For instance, from a 2,000-acre tract of high quality coverts support- ing a fall population of ten grouse to 100 acres or 200 birds in all. 9 per cent or 18 birds might be taken. Were the poj)ulation twice as high (20 to 100 acres) the allowable harvest would approach 50 per cent. One may wonder why it is recommended that hunting be suspended on a good area when the birds are still more plentiful than on a medium area where a take is permitted. But. if a fall population on any area were reduced to the lowest point from which it might be expected to recover, one could expect little or no harvestable surplus the following year. The levels below which hunting is not recommended, therefore, represent populations which should, with average natural losses, produce enough birds the following fall to insure an ade- quate breeding stock. This level is lower for coverts of medium quality since the overwin- tering abilitv of such areas is somewhat less than for better habitats. Of course, the latter could safely be reduced just as low or lower. But to do so would result in reducing the general population level they are capable of supporting. One must also remember that, even though a smaller proportion of the population has been taken when hunting is suspended on the good area, the actual lunnber of birds harvested is usuallv greater. Regulation of Hintinc. More exact regulation of the iiundter of birds taken bv iuinliiig than can be accomplished l>y executive action tiirough sotting o])en seasons and bag limits will rarely be warranted over large areas. On local units under intensive management, however, it may sometimes be desirable to limit the kill to a pred(>tcrmined mnuber based on tlie known harvestable surplus at the beginning of the hunting season*. The need for supplementary control on a managed area implies a very high hunting pressure which, operating freely, might endanger the maintenance of an adequate brood stock. The sim|)lest way of accomplishing this aim is to restrict the number of hunters or the size of their take or both. In order to carry out such limitations one must have complete control of trespass on the area. The land must be posted within the legal requirements of the state law and must be patrolled to prevent poaching. Some system of checking huiitiMs in and out of the area is also needed in order that records of the harvest can be maintained. \\ hen these records show tlial the allowable lake has been attained, hunting of grouse should be stopped for the year. The point may be raised that, over an area, certain good coverts, being in the productive |)hase of their cycle, are well stocked while others, equally good, are i>ut sparsely occupied and should therefore not lie Iiunted as intensively. Theoretically this is true. But in prac- tice over large areas it is, of course, quite impossible to limit the intensity of llic Ininting on * Spc disciiBsinn of Thp I'roinirlion it( tlir Cro|» Available f.'-}). Public interest is best served by producing both timber and game crops with a fair regard for each. (p. 683). The degree of correlation to be sought between grouse management and wood production is a matter of policy and is administrative in nature, (p. 683). In portions of a park, management of its woodlands may well incorporate measures advan- tageous to grouse, (p. 684). The forest preserve type of park, in the Northeast, is all grouse range and is capable of great ■improvement, (p. 684). National Parks present essentially the same problems for grouse management as do forest preserves, (p. 685). Coordinating grouse production on privately owned lands is vastly different than on public lands; a public agency has a responsibility to the people not felt by private owners, (p. 685). r)R2 COORDINATING GROUSE PRODUCTION WITH OTHER PRIMARY LAND USES Iiiterestiii}; landou iicis iji inacticitij; grmisi- inanagemenl cm llicii lands is piiTiiariK a mai- ler of eclutaliiiii. ( |). 685). Un corporate lands, grouse inaiiageineiil ma) Lcsl l)c realized tlirough desirable practices for timber, livestock or crop production as well as for grouse, (p. 686). Catering to the hunting fraternity builds goodwill for corporate owners of large land areas. I p. 686). Soil Conservation Districts and Forest Districts offer an opportunity for landowners to carry out soil, water, forest, and wildlife conservation in a coordinated program with the aid of state and federal conservation agencies, (p. 686). The ruffed grouse is but one of the useful game species produced in most woodlands. Like- wise, game, as a group, is but one of several important natural resources that are derived from the land. It is evident that the degree to which grouse needs may be worked into the handling of an area will depend, to a large extent, upon the primary uses of the land and the ways in which these uses conflict with grouse interests. Since the grouse is a resource of importance to the public at large, its development is of great social importance as well as of interest to landowners. Thus, the type of ownership as well as the primary land use is an important factor in coordinating grouse management with other land use practices. * FITTING GROUSE NEEDS INTO MANAGEMENT OF PUBLIC LANDS Public lands are of many sorts, acquired foi- a wide variety of purposes. The Federal Government owns land in vast acreages for national forests, and in small areas for light- houses, national parks and monuments, prisons, customs houses, and waterfowl refuges, which are just a few of the hundreds of types of use of land owned by the United States. Similarly, the State acquires land for many purposes, some of them paralleling federal uses. Forests, pen- itentiaries, parks, highways, canals and game lands are a few examples. Among the smaller units of government, the county is most likely to own lands extensively. However, other governmental units such as townships and conservatidii districts inan\ also have lands of various sorts under their control. A large proportion of these public lands will not contain important grouse coverts. Off- setting this, those that do include grouse habitats are usually among the more extensive holdings. Of greatest importance are game lands. ])nblic forests and parks. Game Lands There should be little difficulty in meeting grouse cover requirements on most game lands. Where the prin\ary land use is for game, it is merely a question of balancing the re(]uircmcnts of the various species. The conflicts are less likely to be connnon or serious than on otiicr lands. Within this limitation anything the grouse needs may be provided. liniilcd. of course, bv the restrictions of economic feasibility. In the Northeast, the game and fur animals most commonly found on grouse range are cot- FITTING GROUSE NEEDS INTO MANAGEMENT OF PUBLIC LANDS 683 iDiilail rabbit, white-tailed deer, gray squirrel, woodcock, varying hare, raccoon, mourning dove (a game bird only in the south), black duck, beaver, red and gray foxes, New York and small brown weasels, skunk, muskrat and mink. In some places the pheasant overlaps the fringes of grouse range. Only the first six are usually of enough widespread importance to seriously affect land management problems. In their broad aspects, the needs of rabbits, deer, squirrels, woodcock, hares and 'coons are much the same as for grouse. They occupy, in part at least, the same habitats and their cover needs are similar in many respects. Clear-cut plots that are good for grouse are likewise useful to rabbits and deer. Spring nesting grounds are good gray squirrel cover, with due regard for denning facilities. Winter shelter for grouse will serve the same purpose for deer, hares and cottontails. In most ways the one pattern of land use fits the group to a surprising degree. In minor details there must be adjustments, some of which may affect the grouse. If beaver are stocked they may destroy valuable fall and winter food in popple groves. The beaver may also furnish desirable openings by flooding and killing small areas of woodland. These and similar interests must be reconciled in the overall plan. Generally, the broadening of the resource base, as by the addition of beaver, will be warranted for the development of more favorable conditions for native species, even though it may mean a few less grouse. On game lands not in grouse range, or in marginal grouse range, the land use pattern may well be unsuited to the species. Areas mainly adapted to waterfowl and farm game species will offer little opportunity for practical grouse prodiirtion. Here there is no problem of coordination insofar as grouse are concerned. Public Forests The largest public land areas are in this category. Here one often finds real opportu- nities for coordinating grouse production with the primary use of timber production. This comes about not only because of the extent of these areas but because much of this land is in regions where grouse can be produced in significant numbers. Management of public for- ests has given but little recognition to the development of secondary crops, such as game and recreation, until very recent years. Much more progress can be made in this direction. Coordination of grouse requirements with timber growing implies that some modifications in the plans for developing wood products are needed in order to produce on the same land a reasonable crop of grouse. It also implies that the public interest will best be served by producing the two crops with fair regard for each, rather than concentrating on the one alone. This decision is fundamental. The technical methods of correlating grouse management with wood production have already been fully covered.* The problem here is one of policy, and is administrative in nature. It is apparent that the various modifications of forestry practice needed to favor an increase in grouse will only be realized if it is the policy of the agency handling the lands to include these provisions in their various plans and operations. As an example, the planting policy of the New York Conservation Department Division of Lands and Forests on State reforestation areas may be cited. In the early >ears of the acquisition and development of submarginal farm lands for State forests, it was the standard practice to plant all open fields solidly to trees — excepting, of course, lanes needed for fire * Sec Chapter XVI. 684 COORDL\ATL\G GROUSE PRODLLTION WITH OTHER PRIMARY LA\D USES lines. Since these areas were also "Public Hunting Grounds", this policy was of great con- cern to many sportsmen. As a resuh of many conferences between interested foresters and game managers the policy was changed to further both interests. Among the several revi- sions was one to provide that a chain width I 66 feet I be left unplanted on each side of all roads and fencerows. Thus a network of hardwood cover was assured through the new coniferous areas since most natural reproduction would be of that type. Also it provided an opportunity for shrub borders to be planted or open edges maintained in accordance with the grouse or other game needs of the area. The important thing to emphasize here is not only (hat an adjustment in favor of grouse was made but that it was only possible by adjusting the policies of the agency. It is in the top office that the coordination of grouse production with forestry must originate for public ■forests. Parks and Forest Preserves There are two types of state parks in New York and they differ from each other markedly. One is the typical reservation of a unique spot for recreational use, such as bathing beaches, beautiful ravines, picnic groves and the like. In the second class are extensive forest areas that include great opportunities for recreation. The Adirondack and Catskill Forest Pre- serves and the Allegany State Park are examples of the latter. Their management differs from that of a public forest in that they are not managed for the production of wood products. In fact, in the Adirondack and Catskill parks, the State Constitution expressly forbids the har- vest of live trees, which includes a large share of practical forestry, as well as those game management practices involving improvement of the environment by controlling the vegeta- tive cover by cutting. The ordinary state parks are areas of concentrated human activity and hence it has been necessary to prohibit hunting. Even where grouse range exists in a park, this obviously limits the opportunity for correlating grouse production, since the normal objective of pro- duction is harvest by hunting. The cause is not wholly lost, however, for the grouse is also a first class asset to a park just from the natural history point of view. If a park area can support grouse for people to see, the usefulness of the park for recreation is thereby enhanced. Except within the highlv developed (and often artificialized) portions of a park, the man- agement of park woodlands may well incorporate measures advantageous to grouse. \\ hen a woodland border is being landscaped, it can well become a useful shrub border (fall feeding ground) for grouse by a judicious selection of shrub species. When a nature tr;iil is being constructed, it may well provide desirable openings for the grouse. These methods should be worked into the development plan for the area with deliberate consideration of the desired objectives. The maintenance problems must likewise be cared for by specific practices to bo fdlloucd in tiic (ipcr;iti(in of the park. The forest preserve ty|)e of park presents a greater opportunity for |)roducing grouse than parks of the kind just mentioned. In the Northeast, it is all grouse rangi-. usually with a rather low carrying capacity but capable of great improvement. On the other hand, most grouse management in such preserves in New York State is impossible owing to the legal lim- itations on use of the areas. Most of the forest preserve parks are open to hunting, how- ever. COORDINATING GROUSE PRODUCTION WITH PRIVATE LAND MANAGEMENT 685 In wooded land, tree cutting is essential for the development and maintenance of grouse cover values. With cutting of live trees explicitly prohibited, grouse management, in fact most game and timber management, is exceedingly limited in scope and the opportunity for multiple-use of the land is small. The wilderness areas of our National Parks present essentially the same problem in so far as grouse are concerned as do the forest preserves. The objective being to maintain primi- tive conditions as nearly as possible, one cannot expect large grouse populations. Some National Parks offer more opportunity for coordinating grouse production with recreation, especially large forested areas. Here the problem is similar to that of public forests, excepting those portions that are intensively developed and where hunting is not allowed. Most other types of parks, city- and county-owned areas primarily, are small and are usual- ly not suited to producing grouse. Generally they are situated where grouse have long since been excluded by civilization. COORDINATING GROUSE PRODUCTION WITH PRIVATE LAND MANAGEMENT The problem of working grouse improvement measures into the handling of private lands is vastly different than it is with public lands. With puliiic lands, each agency mav handle large areas and, once adequate policies and procedures are established, they become widely effective. A public agency has a responsibility li> the general public that is not felt bv the private owner. Furthermore, a public agency such as a Fish and Game Conunission has a specific responsibility to the sportsmen. Lands Owned by Individuals Today there is very little in our civil laws tliat requires private land owners to use con- servation practices. Each man is King on his own land and he can do about as he wishes, with a few exceptions. He cannot always, in the Northeast at least, burn his woods at will. He must keep his livestock on his own land. He may have to accept certain measures in the pub- lic interest, such as the elimination of gooseberries and currants for blister rust control. But even there, he doesn't have to do it himself. Many of the acts he may commit on his land may be detrimental to grouse. Some may be destructive to his neighbor, or to the general public interest. But except where specific personal damages, not caused by "an act of God", can be proved to have resulted from his negligence, there is no recourse for either his neighbor or the public. He can graze the wood- land, clear-cut it with destructive lumbering practices and otherwise eliminate grouse cover values and no one can say him nay. Socially-minded people will agree this is not as it should be. Some day there may come to pass a change whereby conservation ordinances will prevent such destruction. But until there are greater limitations in the use of land and its resources by the individual, grouse and other game production will be either accidental or the result of personal interest. The development of the interest of the landowner in grouse production to the end that he will practice grouse management on his lands is primarily a matter of education. If he is interested in hunting, he may be a willing student. If he just likes to have grouse around he may likewise do many things in their favor. If he has no special interest in having more birds, the only way that beneficial practices can be coordinated into his operations is by indirection. He may benefit grouse through means aimed at some other objective. Ul',6 COORDINATING GROUSE PRODUCTION WITH OTHER PRIMARY LAND USES When livestock are fenced out of a woodland as a measure in animal husbandry, grouse may be benefitted just as much as though it were done delilteralcly for their ititurests. Shrub borders that are planted (or developed by cutting) and maintained for woodland protection, for crop field protection, or simply for their beauty may be just as useful to grouse as though they were planned for them. By educating farmers and woodland owners to good land man- agement practices, much improvement of grouse cover may be accomplished. Lands Owned by' Corporations Private land has been discussed as a problem of individual ownership. Much of this land today, especially large forest lands, is owned by corporations or other similar groups. Here one cannot a])i)eal very strongly to personal interests for the land to them is an impersonal matter. Things which are not good business, that is, economically profitable, are not likely to demand their attention. Since grouse are a somewhat undependable crop at best, and in any case are owned by the State and may not be connnercialized, grouse management in itself has not to date been considered a profitable enterprise. Attaining grouse production on corporate lands may best be developed through practices that are desirable for either timber, livestock or crop production as well as for grouse. Over and above this group of agricultural and forestry operations, some modifications in favor of grouse may be promoted through the public spiritcdness of the organization. More and more companies are realizing the value of good public relations and those operating large land areas should certainly find adequate reason for catering to the hunting fraternity. Soil Conservation Districts The history of conservation endeavors on private lands shows clearly the need of technical assistance on the ground if owners are to become practicing conservationists. The coimnon educational methods are not enough. Formal schooling, written articles, bulletins and books, public demonstrations, radio talks, and the like, invaluable as they are, have not succeeded in stemming the tide of destruction of our natural resources the land and its products. Ad- ditional assistance to private owners has been provided in a number of ways in recent years. The furnishing of tree planting stock for reforestation is a good example. These seedlings are grown by state nurseries and sold to tlie public for a very nominal fee to encourage proper use of lands not suited to growing farm field crops. Farm foresters and district game managers provide technical advice tailored to the specific needs of each property so that the planting stock may be most effectively used. The growth of the national soil conservation program has helped to clarify some of the principles that are essential for successful conservation work on private lands. Four of the cardinal points are: fl) conservation ]>lamiing should be based on an accurate physical in- ventory of the land. By mapping the area according to its inherent capabilities, the proper use can be determined; (2) the various sciences must be coordinated to produce best re- sults. As an example, the needs for growing wood jnoducts and wildlife in the same wood- land require the harmonious fitting together of the sciences of forestry and wildlife manage- ment; (3) the complete land ownership enterprise should in- |ilanned as a unit. All parts of a farm are interrelated as portions of a single business. Only by planning the entire farm at once can the proper land use and needed conservation practices be assured; (4) relations among private landowners and conservation agencies must be democratic. This principle of a democratic approach to the conservation job on jirivate lands de- COORDINATING GROUSE PRODUCTION WITH PRIVATE LAND MANAGEMENT 687 serves special attention. Fortunately, it has received a lot of thought in the past decade. In 1937 a number of states passed laws enabling the establishment of a new political subdivi- sion called ihe Soil Conservation District. Now all 48 slates have such a law. They vary somewhat in their specific provisions but all have the same objective — to permit landowners to obtain help on conservation problems in an efficient, democratic way. New York's law was passed in 1940. The philosophy of the soil conservation district is to apply grass-roots democracy to con- servation work. In the beginning, the stimulus for the creation of a conservation district in any area comes from the people. Through petition, or by personal appearances before the county board of supervisors, they make known their desires. After adequate considera- tion by public hearings, the county supervisors decide by vote whether the county shall be- come a soil conservation district. (In many states, a favorable referendum by ballot by the landowners is required). During the period when interest in such a conservation district is developing, the Exten- sion Service of the State College of Agriculture takes the leadership in a program of infor- mation and education. Later, after the district is created, this educational program is con- tinued to encourage landowner* to seek aid in conservation work. The directors of the district are appointed from among progressive, local farmers. They begin the work of the newly- formed district by developing a ])rogram setting forth the conditions that make conservation work urgent and the objectives of the district. Then a work plan is prepared to set forth the methods by which the district will operate, including the types of help that will be pro- vided by the different cooperating agencies. Experts from these agencies are ordinarily called in for advice in preparing these documents. When the organization is thus completed, the men, materials and equipment facilities are ready for business. Landowners may apply to the Directors for on-the-farm assistance in the planning and establishment of a complete conser- vation plan. It makes little difference what conservation problems an individual land owner has; they all fit into a sound plan for soil conservation. Thus, grouse management practices go hand in hand with the control of gully erosion, contour strip-cropping, and field drainage. The owner applies for aid to tlie directors of the soil conservation district in which he lives. The facili- ties of the district, made available through its cooperating local, state and federal agencies, are applied to his farm on a voluntary basis. A conservation survey provides the needed physical inventory. Based on this survey, and on the economic needs of the farm business, a com- plete farm conservation plan is developed by a planning technician. This service is gener- ally made available to the district by the U. S. Soil Conservation Service which is especially set up to carry out this function. Special services to implement the woodland and wildlife phases of the conservation plan may be rendered by technicians from the forestry and game divisions of the State Conservation Department. Thus, the available conservation services are funneled to the landowner through the facilitating soil conservation district. Forest Districts A recent development that should become of great importance in grouse management in New York was the enactment in 194G of the Forest Practice Standards Act. Its purpose is to provide a voluntary, cooperative program whereby over-cutting of woodlands may be avoided, forest product industries stabilized, and sound forestry encouraged. It provides for the es- tablishment of forest districts, district forest practice boards to determine standards needed 688 COORDISATING GROUSE PRODI CTIO\ WITH OTHER PRIMARY LAM) USES in forest practices, and a state forest practice board to review district forest practice standards and promote the welfare of these forest districts. The chairman of each soil conservation district within the forest district is autoinaticallv an ex-oflficio member of the forest prac- tice board to assure integration of the soil conscr\ati()n district program with the forest prac- tice standards developed for the area. The State has been divided into 15 forest districts. Each has its own board consisting of three members from each county included who are appointed by the chairmen of the re- spective county boards of supervisors. Only one of these three may be chosen from the county board of supervisors. At least two must be resident woodland owners. The state board is made up of the chairmen of the 15 district boards plus ex-officio members, without vote, from several state agencies. These district boards determine the forest practice standards necessary for their districts. After adoption, these standards are promoted h\ the district on a voluntarv basis. However, woodland owners who desire the technical forestry services furnished by the district must agree to abide by these standards as a condition to the furnishing of the aid. It is anticipated that these standards will be fairly simi)lc and will have as their basis the control of fire and grazing and the marking of harvest trees by a forester prior to lumber- ing. This would assure adequate protection of these woodlands and the use of a cutting plan that would assure future productivity of wood products. These sorts of standards fit well into the needs of grouse management. It is conceivable that the practice standards might go even further in areas of good grouse range and include more detailed cutting practices that would be definitely aimed at improving grouse cover. But that is still in the future. We must wait and sec how it develops. The significance of the soil conservatinn and forest district developments lies in the fact that they are symbols of a new conservation era. They are tools of societv. de\eloped to make conservation of these resources a realit) through a democratic svsteni. They repre- sent substantial progress toward genuine working cooperatimi b('lw<'(ii tlic \arious interests concerned. They facilitate the inclusion of grouse management to an\ desired degree. By the same token in other situations also it will be only through sincere, mutually coop- erative elfort. both in planning and in action, that effective coordination can be achieved. Lack of such team work will result either in ineffectiveness or wasteful (lii|ili(;ilion. "Ding" Darling once said that ten thousand wild horses free on the range conldn t pull a rubber- tired baby carriage. But harness those to pull together and they can move '"mountains"'. SoLKCKs OF Technical Aid The need for technical assistance in coordinating grouse nuinagement with other land uses has been noted. In New ^ mk the district game managers of ihc Conservation Department represent the primary source of such aid. Advice on special problems is available from wild- life research |)ersonnel. district foresters and forest technicians of the same Deparlnicnt. and from the Kxicnsion Scr\ ice of the State College of Agriculture. In connection willi soil con- servalion (listricls in pailic uiar the U. S. Soil Conservation Service is importanl. \ ahiable infornuttion can also In' obtained from the U. S. Fish and \\ ildlife Service. The fact that such assistance is available, however, will result in no benefit to grouse or other wildlife unless it is actual!\ jiul to use on the land. THE FUTURE 689 THE FUTURE The opportunities for integrating grouse management with other land use interests have been discussed earlier in this chapter. Similar possibilities exist for other forms of wild- life. Means of implementing such coordination are being developed. It is up to us to make them work. The alternatives do not make a pretty picture. Either we manage our soils, tim- ber and wildlife on a sustained yield basis through voluntary methods in a democracy, or we waste them to the point where we become a second-rate "have-not" nation, or we force restrictions on their use by government control, autocracy, communism, or whatever else you may wish to call it. Which will we choose? '^::::--i ....«,. '^ yi^?^\ IN MEMORIAM 691 IN MEMORIAM As we idly finger the pages of this book, our work seemingly done, we realize with a guilty start that, in the very last sentences we have left the poor grouse fluttering in the dubious care of the communists. Surely, this is no fitting end for such a gnllant friend, the source of years of research, work and fun. It is our desire, as well as our duty, to make the parting fit the honor we owe him. The Investigation has been extensive. We have gone back over twenty-five thousand years to dig him up. We've played with him among the Indians and pioneers: slaughtered him with sticks and guns, and trapped and sold him to greedy markets. We've invaded and de- stroyed his home and designed new mansions for him; sampled his food and prepared new menus; robbed his nests, adopted his babies and nursed them for better or for worse. We've tried to propagate and manage him, practiced eugenics on him, psychoanalyzed him. probed his insides, studied his diseases and exposed his parasites. We've consorted with his enemies, we've swapped stories and spun yarns about him. We've thrilled to his quirk wil and trick\ flight; stood s])el]bound listening to his drum- ming and watching his love antics. We've discovered he's a healthy specimen and lost most of our preconceived notions about cycles and the grouse disease and pet fallacies of what causes the fluctuations in his abundance. He's still a mystery, still unpredictable and, be- cause of our research, a bigger, broader problem than ever before. With it all, we love him for what he has been in the past, what he is in the present and what we hope he will be in the future. He may have thrown us for many a technical loss, he is a stubborn subject, but science refuses to be discouraged — the work will go on. No re|>ort or book will ever write finis to a scientific subject as complex as is the grouse. Keen minds will conlimie to delve ever deeper into this fascinating riddle. To them, and to you, oh reader, who has waded with us through this labyrinth, our deepest sympathies — and to the ruffed grouse our humblest regards. The Authors. I t PART IV Appendix METHODS AND TECHNIQUES By Walter F. Crissey ORGANIZATION OF THE INVESTIGATION GENERAL METHODS Selection of Study Areas — Mapping — Personnel — Securing Observations — Recording Observations SPECIAL TECHNIQUES Determining Shelter Relationships — Determining Food Relationships — Deter- mining Weather Relationships — Determining Predator and Buffer Relationships — Predator Abundance and Activity — Buffer Abundance — Food Habits of Predators — Predators Responsible for Grouse Kills and Nest Destruction — Determining Effect OF Hunting — Determining Disease Relationships — Estimating Grouse Popula- TioNs — Trapping and Marking Grouse — Determining Sex and Ace USE OF BIOMETRICAL ANALYSIS Significance — Chi-Square — Analysis of Variance — Regression and Correlation SS The success of any project is greatly influenced by the methods and techniques employed. At the beginning of the Investigation there was a lack of standard means for carrying on such work since most of the ])ioncering, u])on which cdicient methods were eventually based, had only just begun. It was natural, therefore, that the process of standardizing methods, weeding out unsatisfactory techniques and the production of usable knowledge should have gone hand iti hand during the course of the study. Roth sportsmen and wildlife research workers are naturally interested in knowing how the records were collected and analyzed as well as what conclusions were drawn from them. To that end, it is the purpose of this section to discuss briefly the organization of the Investigation and. in more detail, the methods which were newly developed or adapted from existing wildlife research techniques. ORGANIZATION OF THE INVESTIGATION The Ruffed Grouse Investigation was the direct result of the great scarcity of grouse in New York State in 1927-28. At the request of a special committee of alarmed sportsmen* funds were provided by the 1928-29 Legislature for a thorough study to determine and eval- uate as many as possible of the causes underlying the varying abundance of ruffed grouse * See Chapter I. 694 METHODS l\n TECHMQUES and. if ])ossible. to find and develop means of producing and maintaining; high grouse ])i>[)iiIations. The Investigation was organized and has heen directed throughout by the senior author Gardiner Bump. Field work began in March 1930 and was carried on continuously until September 1942. Artificial j)ropagation studies and laboratory investigations of disease and parasitism were expanded with the establishment of the New York State Game Research Center in 1933. The analysis of data secured was completed in 1943 as was the first draft of this report. Publication was. however, delayed until 1947 by the war. The first step was to outline the problem and plan the work so as to secure the maximum information available. The work was organized around the following ten major factors which affect the production of grouse: 1 . Physiology 4. General Habits 7. Parasitism and Disease 2. Shelter 5. Weather 8. Alan 3. Food 6. Predators 9. Rejiroductive Capacity 10. Artificial Propagation (Replacement of seed stock) These were in turn subdivided into 253 potential problems. The work was conducted and summarized largely around this original outline. Study areas were selected and tentative survey methods chosen to collect as much data as possible. As the information accumulated, the important problems became more clearly de- fined. Techniques were checked and altered, where inadequate, to insure evidence that better represented the existing conditions and could be analyzed for valid conclusions. As the study .progressed and expanded it became evident that no one person could be responsible for all the specialized details. The res|)onsibilitv for carrying on most of the field work was accordingly divided between the two jinu'or authors Robert \^'. Darrow and Frank C. Fdminster. Jr. The latter left the Investigation in 1937 and was rephtced liv Walter F. Crissey. The following s|)ecia]ists were engaged either to direct work on s|iccialized prob lems or to collaborate with the authors in their solution: food habits. .Tohn C. Jones; para sitism and disease. Dr. P. P. I.evine and Dr. Fraiis C. Goble: artificial |iro])agation. Dr. A. A Allen, Earl R. Holm and Janel W. r!um]i: physiology. Dr. Wm. H. Long: embryology. Dr, Alexis L. Romanoff; pterylography. John E. Trainer; anatomy. Dr. David E. Davis; biomet rical analysis of data. Dr. Masoti I-awrence and Dr. C. McC. Mottley: and bibliography. Dr Mary Thornton and Albert G. Hall. GENERA!, METHODS In setting up the Investigation there were certain general considerations which preceded the actual work on specific problems. They included selection and mapping of representative study areas, choosing of personnel, as well ns deciding on the methods to be used in secur- ing and recording data. Selection of Study Areas Considering the .Stale as a whole, three major regions were recognized on the basis of habitat difTerences. The Adirondacks are distinct in comprising extensive forest areas of a largely coniferous climax tyi)e and having a more severe winter climate: the Catskills are a rugged region consisting of a heavily forested nn( Icus supporting a more |)n'dominantly hardwood cover, surrounded b\ a zone where considerable agricultural develupmcnt has taken place; the "Rest of the Stall-", typified by the central and Southern Tier ])lateau, is character- GENERAL METHODS 695 ized by disconnected woodlots, primarily hardwood in composition. Accordingly, four pri- mary study units*, one each in the Adirondacks and Catskills and two (the Connecticut Hill and Pharsalia areas) in the plateau region, were selected in order to facihtate recognition of any differences in relationships which might exist. Also, from time to time, special problems have been studied in other areas. The following important items were among those consid- ered in choosing these areas: 1. Representativeness — whether typical of the region in which situated 2. Size — sufficient to yield adequate data 3. Control — public ownership if possible The major study area, Connecticut Hill, is located in the plateau section of southwestern Tompkins County, overlapping slightly into Schuyler County. The basic portion comprises some 2,200 acres of grouse cover with many intervening tracts of open land. Formerly an agricultural district, the farms have been largely abandoned during the past 30 years. In consequence, the fields surrounding the woodlots are becoming overgrown with bushes and small trees. The woodlands exhibit a high degree of diversification and, in spite of having been extensively cut over for many years, contain a considerable admixture of conifers, chiefly hemlock. The topography is shown in figure 71, and the relationship of wooded and overgrown land to open land in figure 72. The cover type composition of a typical compart- ment is illustrated in figure 14 ( |). 17! I. The Pharsalia area of some 2,100 acres lies in northrentral Chenango County along the divide between the Chenango and Otselic valleys. It is quite similar to Coimecticut Hill but the woodlands are somewhat more extensive and contain a number of wooded swamps. The Catskill area of some 850 acres is situated along the northern margin of the Catskill Region in southern Albany County. Its woodlands are also broken up bv intervening fields, but the latter are relatively less overgrown because most of the farms in the vicinity are still operated. The Adirondack area is a forest tract of slightly over 800 acres lying in an upland valley on the eastern slope of the Jay Mountain ridge in northeastern Essex County. A few small clearings along a road which ])artiallv bounds the area constitute the only open land present. Likewise the only overgrown land is found in conjunction with these clearings. The cover is mixed second-growth timber, conifers being chiefly s|)ruce and balsam. The area also con- tains two large alder beds. The topography of this area is shown in figure 71 and its cover type composition in figure 15 (p. 173). Mapping The construction of adequate maps of the study areas constituted the next step preliminary to conducting field surveys. Starting from an existing base, details were usually filled in by the "pace and compass" method although "tape and compass" lines were often used for greater accuracy. United States Geological Survey maps formed the base maps in 1930. but revisions and additions were made from aerial photographs as they became available. The degree of detail was controlled by the requirements of the research to be conducted. Wliere mere reconnaissance surveys were adequate, details were only roughly drawn in. Where the actual location and amount of each cover type were desired they were mapped * Set figure 13, p. IH. 696 METHODS AND TECHNIQUES A P) I MILE CONHECTCUT HILL STUDY AREA AOIROHOACX STUDY AREA FICURK 71. TOPOr.RM'in OK CONNECTICl T 1111.1. AND ADIRONDACK STIDY AREAS GENERAL METHODS 697 liiilliil Wooded 8 Overgown Land FIGURE 72. RELATIONSHIP OF WOODED AND OVERGROWN LAND TO OPEN LAND ON CONNECTICUT HILL 698 METHODS AND TECHNIQUES with greater care and accuracy. In the beginning it was thought sufficient to map specific cover units down to one acre in extent. Further work indicated, however, that units much less than an acre in extent influence the distribution of grouse, and the areas were accordingly remapped showing cover types to one-tenth of an acre. Since records were desired, not only of the type of cover but also of such items as food and shelter and cover density, a fractional system of combining symbols describing these items was developed*. That is, the cover type designation'^ was shown as the numerator with numerals representing cover density, food conditions and shelter value, in that order, as the denominator. Cover density for each ty|)e was indicated bv a classification running- from 1 to 10, 1 representing minimum and 10 maximum density. Thus an open grassy area, providing the grass was tall and thick, was given a density rating of 9 or 10. A woodland, on the other hand, might be classed as 3 or 4, if the crown cover were relatively open. Food conditions for grouse, the second in the series of three symbols making up the denominator of the fraction, were indicated by numerals from 1 to 5. The following differen- tiations were recognized: 1. Poor during all seasons 2. Adequate or abundant during seasons when type was used by grouse 3. Fairly adequate during all seasons 4. Adequate throughout all seasons 5. Abundant throughout all seasons The analysis of shelter value was symbolized by the numerals 1 to C, again interpreted from the point of view of the grouse. 1. Adequate shelter absent 2. Adequate shelter scarce 3. Shelter conditions superior to 2, but not yet adequate throughout all seasons of the year 4. Adequate 5. Abundant 6. Dominant to the exclusion of most food species. In extensive units of a single type, the fractional symbol rejirpscnting existing conditions of food, shelter and density was written in the proper location as the food, shelter or density in that particular portion of the type changed. As an example of the use of the above system, a fairly dense piece of second growth wood- land composed of beech, birch, hop-hornbeam, maple and a few scattered hemlock (food fairly adequate all seasons and shelter adequate) uoiild be described as follows : -r-r-r . A 20-year-old jiiiic plantation, on the other hand, w(iul, N. Y. S. Cona. Dcjil.. Btiroaii of Caino, null. 1, 190 (mini. eographed) . A Scd Chapter HI. t Excrpl the Adirondack area wliirli is rtmfnliaUy solid (orral. GENERAL METHODS 699 compartments of a size that could be conveniently covered in one day by a small crew of men. The compartments varied from 130 to 300 acres of woodland and as far as possible each was surrounded by open land. Some blocks of woodland were too large and blazed lines had to suflSce as boundaries. These sub-units of the larger areas were mimeographed for field and office use. The division into compartments also aided in the analysis of data by providing a means of obtaining sampling-variations. Observations regarding a single item were repeated for each compartment and the consistency or degree of sampling error could be measured for each year. For example, grouse flushes in relation to cover type when analyzed by compartments within years, yielded a much better picture of the consistency of the grouse cover type use relation- ship. Compartment maps were made to a scale of 660 feet to the inch. These were mimeographed in outline form on 8V2 by 14-inch paper with proper titles. In addition to field and office use, the more detailed maps were employed in many prob- lems dealing with the extent and interspersion of various cover types. Personnel Great care was taken in selecting field men. To this end graduate or undergraduate college students with training in wildlife management or in closely related fields such as forestry, botany or zoology, were employed where practical. The field forces were made up of the authors together \silh leaders and assistants. The authors supervised the work on entire study areas, while crews consisting of a leader and his assistants were responsible for the various compartments within an area. The leaders, chosen for their experience or ability, recorded the data. The conscientiousness of the leader was an important item affecting the results, since the assistants were mainly observers. It was found that less care was needed in selecting the latter grouj). Relatively untrained men could be initiated, in a short time, into the more or less standardized method of covering the compart- ments and observing data. College students, older boys from Scout groups, CCC boys, WPA men and local residents have, at different times been used as assistants. In connection with the work of analyzing the field records, draftsmen, bibliographers, stenographers and clerks were employed. To some extent this force consisted of hand-picked labor from WPA and student NYA programs, thus reducing costs. A survey of this kind cannot be carried out by field men alone. Therefore, special field and laboratory work, requiring technically trained men, was handled by ornithologists, mamma- logists, entomologists, botanists, foresters, engineers, nutritionists, biochemists, physicists, statisticians, trappers, food habits investigators, physiologists and pathologists. Securing Observations The scope of the problems confronting the Investigation was very wide. Therefore a com- plete (100 per cent) coverage of each study area by the strip method was used to yield the most satisfactory data pertaining to the greatest number of the proposed problems. To carry out this method crews consisting of leaders, each with from one to four assistants, according to tlie size of their compartments, were used. The men walked abreast at a given distance apart, thus covering the unit in strips, back and forth from one edge to the other. The end men acted as guides utilizing land marks or a compass to maintain a reasonably straight line. Since cover utilized by the birds in relation to time of day constituted one of 700 METHODS A.\D TECHNIQUES the problems, the working pattern of a compartment was changed on successive days. The intensity of work varied with the objectives and the season. Durinor the fall and winter, when the woods were relatively open, a distance of 60 to 80 feet between men proved satis- factory. Distances of 30 to 50 feet were used for spring population estimates and summer GROUSE SURVEY FIELD CKEW LIiNEI) LI' KEAUV TO ENTER A COVERT ON THE CONNECTICUT HILL AREA brood studies, while a reduction to 20 feet between men was necessary before most of the grouse nests could be located. On the average, a crew of two men working 50 feet apart was able to cover about 150 acres ])er 8-hour day. During the first two years of survey, dops were used by some of the crews. It became apparent, however, that they often caused confusion and tended to introduce variables that were hard to account for. Their use was, therefore, discontinued in fa\(ir of the MKire easily standardized crew of assistants. Recording Observations At the outset it was recognized that only by standardizing the method of securing and recording information could its significance be accurately apjiraised. Continuity of the method by which notes were taken was of prime importance in determining yearly trends in relationships. If the method were changed and the effect not recognized, a fluctuation in some factor ])ertaining to a problem would have seemed to exist when, in actuality, it did not. Standardization was therefore the keynote followed in all note-taking. Since many indi- viduals were involved in the field work, the first ste|) was to design uniform record sheets around the more important f)r complicated jiroblems. New compartment leaders were trained and given explicit written instructions on how to record observations. Notes were frequently checked for continuity of handling. SPECIAL TECHNIQUES 701 Note-taking was also designed in such a way as to eliminate, to the greatest possible extent, personal judgment on the part of the compartment leaders. When a personal interpretation, such as the determination of a nest predator, was necessary, the leader referred the matter to the director of the survey. Thus the handling of such decisions rested with a relatively few persons. The coippartment crews were used mainly to collect the data and not to interpret them. Records were taken in such a manner that a daily numerical summary of the basic material could be kept. The information used in reducing data to a common basis, such as weather and man hours, was carefully recorded on each set of notes. Daily notes were then com- bined into seasonal totals, reduced to a comparable basis with other seasons or years and then analyzed. The amount and character of the information gathered was so closely determined by the organization of the data sheets that they are included here. It should be noted that code spaces are provided on each sheet to facilitate the mechanical tabulation of the data (.figures 73 to 76). SPECIAL TECHNIQUES Gathering evidence pertinent to the specific problems involved, often required special tech- niques. Those used in collecting data to evaluate rclatinnships of shelter, food, weather, disease or predator and buffer species to grouse, all differed from each other in many respects. Determining Shelter Relationships A man usually sees a forest in terms of firewood or lumber whereas it is im|)()rtant to a grouse in terms of its food and shelter value. The first step, therefore, was to subdivide grouse habitats into types according to their food and shelter value to the grouse. Since many individuals were to record shelter data, the subdivisions formulated were reduced to letter symbols*, described in brief form and mimeographed for use on a reference sheet. A grouse uses a wide variety of cover types, perhaps more of one than another. When a survey crew flushed a bird, a record was made of the cover it was in. However, there was no indication of the relationship of any one such flush location to the cover requirements of the particular bird involved or the species as a whole. In other words, data regarding the preference of various cover types was extremely variable in nature and immense numbers of records were necessary before any real understanding of the relative degree of use or of the underlying reasons therefor could be obtained. A satisfactory number of records was built up by filling out a data sheet each time an adult or a brood of grouse was flushed during the entire period of survey. Over 19,000 adult and 1,500 brood flushes were thus recorded by 1937 when their tabulation was undertaken. In order to handle this data efficiently, provisions were made for mechanical coding and sorting. Each individual item on the data sheets was given a descriptive code number and these were punched on code cards. After a specific problem was set up. it was possible to run the cards through a mechanical "sorter" and thus obtain the totals of the items desired. Shelter relationships were analyzed around the statistical principle of deviation from the random or "no choice"^ distribution of the bird. Thus an establishment of the pattern in .» Sei- ChapUT HI. A If a {;rinise exercises no choice of the type of cover it uses, flushes should be evenly distribuleil tliroughuut the area and flushes within a single type would occur in proportion to the extent of that type. 702 METHODS A>:D TECHMQiES Weather: (encircle correct condition) a. warm, normal, cold h. sunny, cloudy, rainy, snowing c. strong wind, gusty, still Ground: dry, damp, wet, "snow. GROUSE FLUSHED Serial luirnln'r Date Survey Compartment . Names Type of data Data Code Data Code Data Code 1. Map number 2. Direction you are going 3. Direction flushed (N, E, S, or W) •1. Distance from bird (.in feet) 5. Time of day 6. No. flushed together 7. Sex(es) (J , 5 , or ? Based on: fl, plu, dr, nest, artificial mark. 8. General cover 11. Slope: 1. Compass dir. down 2. Flat, gentle or sleep b. Type of cover 9. Exact flush location a. From the ground b. From a tree (species) 10. Mclluslwd (yes or no) a. Type of cover b. Map number c. Direclioii rrlluslicd 11. Factors inllucnciiig position of liird 12. Ilemarks: (if long, refer to notes) Fir.lRF. 73. DATA SHEET USED TO RECORD INFORMATION RECARDINC. ADI I.T f.ROt SE FI.tSIIEn SPECIAL TECHNIQUES 703 GROUSE NEST County Serial number Locality Survey Year . Compartment, Name 1. Map No ; 2. Nest No ; 3. Date found 4. No. eggs: a. When found ; 1). Total clutch 5. First nest or renest 6. Female flushed: yes, no. a. Dist. from observer ft. b. Direction flushed ; c. Distance flown ft. <1. Bird feigned injury: yes, no. 7. Number of enlarged droppings found a. Dist. from nest ft.; b. Direction from nesi c. Downhill, level, uphill from nest 8. Nearest dr. log: DL No ; Dist ft.; Direct, from nest 9. Distance from: woodsroad or trail, public road ft. 10. General conditions prevailing in nest vic'inity: a. Slopi': Direction (downhill) ; Degree: (lat. gentle, steep ; b. Hcliition to nearest opening: Distance from fl. Type of opening c. Nearest slashing: Size acres; Dist. from ft. d. Type of cover c. ( Jrfmiid cover type f. Undergrowth: Type Specie's ; Density g. Crown cover: Species Density 11. Specific site of nest : n. l?esidc or in: tree. stum|i. brush pile, rock, log, clump of bushes, cavity in stump or bank. op<'n forest floor, open field b. Species (if at base of tree) c. Diameter 1 foot from ground (if at base of tree) d. l">\posurc (diri'ction it faces) e. Distance from nearest conifers ft. f. Density iinmeilialc situation: in open spot, in thicket IvXrh: OF M'lST 12. Hatched: n. Number of eggs hatched ; Dati! 1). Number left unhatclied: No. infcrtili? No. partially developed c. Number not known whether hatclicd or not d. Condition of hatched shells: No. left as hatched ; No. partly broken WCre any broken to bits: yes, no; No. missing \\ Cre any shells out of nest: yes, no; Dist ft. e. Had nest evidently been disturbed aiftcr hatching; yes, no. 111. Broken up: a. Date b. Condition of nest c. No. and condition of shells found d. Destroyed by: based on : e. Quality of evidence: excellent, good, fair II. Taken for propagation 15. Evidence inconclusive Remarks: 16. Date checked ; By FIGURE 74, DATA SHEET USED TO RECORD GROUSE NEST INFORMATION 704 METHODS AND TECHNIQUES GROUSE BROOD FLUSHED Weather: (encircle correct condition) Serial number. . a. warm, normal, cold Date I), sunny, cloudy, rainy, snowing. . Survey c. strong wind, gusty, still Compartment.. (iroirnil: dry, danip. wot "snow Name Brood number. 1 . Map No ; 2. Time of day ; 3. Contact No 4. Distance from: a. Female ; b. Nearest chick 5. .\mounl of lime spent with brood minutes. 6. l)irc<:tion you are {;oin},' 7. Direction: a. Female flushed ; b. Chicks flushed 8. General cover from which (lushed (answer a, b, e and d): a. Slope: 1. Compass direction down 2. Flat, gentle, steep b. Distance from opening ft. to type (Use .\,B,C,D,I,J,\VR) c. Type of cover (Use A,B,C,D,E,F,G,E-H,F-H,I,J) d. Distance from crown cover (if flushed in open) feel to type 9. Immediately surrounding cover (within 100 fc^et of flush): a. ('rown cover (''o o{ types represented) b. Undergrowth: 1. Type ; 2. Density c. (iround cover: I. Type ; 2. Density 10. Exact cover fniin which flushed (within 10 feet of (lush |)oint): a. From the ground (yes or no), .\nswer 1 or 2 and 3. 1. Close to trees or shrubs; (a) .Specie-s ; (b) Height 2. la the open (at least 10 feet from trees or .shrubs): (a) Distaiure from trees or slirubs ft. (b) Species of trees or shrubs 3. E.xact description of species of cover at jMjint of (lush: (a) Urowii cover: species density. (b) Undergrowtli: species density. 1). From a tree: species ; tree height ft.; bird height. . . .ft. 1 1. Male grouse flushed nearby (yes or no) ; distjmce from brood 12. C(mdit ion of brood: a. Nundier of chicks: 1. seen ; 2. hi'ard, not seen b. Kstiiiialed ninnber of chicks in brood c. Size of chicks in cdniparisoti with a robin d. \ge of brood, eslinialed in weeks e. Brood bunched or scattered when Ihishi^d. . ; how far f. Developmt^nl of plumage 13. Behavior and movements of female, of brood and of men on .survey 1 I. Ideiilily of brood baseil on: 1.3. Probable influencing factors: a. Regarding po.sition of brood : 1. b. Regarding behavior of brood : 1 2 c. Regarding methods of discovery and handling: 1. 16. Quality of brood contact us to nundicr of chicks -. 17. Markings of female: a. \rti(icially marked or not (yes, no or ?) Ii. If tiiarkeil. giM' color c. ( 'olor phase of female 1({. Remiirks: (if long, gi\e reference to position in notes). H(;LRK 75. D.\T.\ SHEET USED TO RECOKP (.KOI SK BROOD I.NFORMATIO.N SPECIAL TECHNIQUES DEAD GROUSE FOUND 705 Original Kill. Other Parts. Serial number. Date Survey Compartment. Name 1. Map nuniher ; 2. .\dult or chick 3. Sex ; Based on 'I. Color phase ; I^eiigth of t.'iil feathers 5. Field data: a. Cover: 1. In an open area (.\ or I) or wooilsroad Distance from woods or bnishland ft. 2. In cover type Distance from open area (A or I) or woodsroad ft. Type of open area or woodsroad b. Number of points where remains were found, their size and distance apart c. On, under or near what found. d. Predator sign found with kill. . e. Wen; any portions of the bird found "in situ"? f. Age of remains (in days, weeks, or months) . . . Laboratory data: a. Quantity of reniains found: liones: Feathers: tail ; wing b. Evidence found on feathers: contour. c. Eviilence found on bones : Tail Pri- maries Other Wing Feathers ( '.on- tours Number Niiiiilier beak- niarked Distance between Number clii>ped over 1" above base. . Number clipped at base 7. Cause of death 8. Quality of evidence: a. Exc(^llenl, good, fair. b. Based on 9. Remarks: FIGURE 76, D.\TA SHEET USED TO RECORD INFORMATION REGARDING DEAD GROUSE FOUND TOf. METHODS, ,1\D TECHMQIFS which flushes would occur in relation to the various cover tvpes. if grouse exercised no choice of them, was a necessary preliminary step. For the purpose of the ln\cstif.'ati(iii. the ''no choice" pattern, witli one exception, was set up on the hasis of the amount of ear li t\ pe ])resent. Three of the studv areas were located in a haliitat t\pe characterized 1)\ ahaiuloned farm.*, so the woodland, though sizeable, was surrounded by open fields. Since these are not used by grouse except along the edges border- ing the woods, an arl)itrar\ limit of TOO feet from existing wood or brushland was taken as the extent of open land that should l)e included in figuring the amount of each type present. It was also ob\'ious that the habit of frequenting the edges, rather than the centers of large blocks of cover, excluded other areas from the birds' use. Therefore, a measure of the amount of each type available was better than the total amount for determining the pattern of occurrence which would be associated with no choice regarding cover type. No practical method existed, however, for evaluating availability. The reliability of the information was checked by two methods. If 20 per cent of an area was composed of a certain type of cover and 25 per cent of the flushes during a ])eriod occurred in that type, the validity of the difference was determined by the statistical method of chi-square*. More detailed problems, such as those dealing with difference in cover type usage between areas, seasons or years, were checked by analysis of variance. Determining Food Relationships The items of food upon which the grouse subsists were determined by an examination of the gizzard and crop. The birds collected during the course of the Investigation were taken by various means and by a variety of individuals. Satisfactory sampling of the grouse popu- lation has de|)ended on the cooperation of sportsmen, naturalists, game |)rotectors. special collectors and the public in general. An attemjU was made to maintain a uniform flow of birds to the Research Center in order that continuous infortnation by regions during all seasons throughout the period of studv. with a good re|)resentation of both sexes and different age groups would be |)rovided. Through the cooperation of the above-mentioned groups, birds have been received which were found dead in the field, hit bv automobiles and trains, those which have flown into houses or wires, and others killed by predators a.s well as a considerable number taken during the regular open season. The backbone of the continuous study, however, was formed by collections made by shooting specified numbers of birds from designated areas at pre- scribed times. For this purpose, special collectors were hired as needed. Certain game protectors were also assigned to the job. The standard methods of procedure as developed and described"*' "^ by the Section of Food Habits of the U. S. Fish and Wildlife Service, were followed in making the analvses. Food items difficult to identify were referred to specialists. Simple but important refinements in the method of determining the average volume of each food item, as obtained from a number of analyses, were adopted. Previously, the volumetric percentage of a single item in the total diet has been figured by first finding its percentage in each individual bird, then averaging these to secure the jjercentages for all birds. This procedure is, of course, mathematicallv unsound. For example, one bird may eat only two beechnuts, rejjresenting 100 per cent, while in another, the same tiumber makes up oidy 10 per cent of the volume. Averaging these, four beechnuts become rin |)er cent of the total volume. Obviously this is not true. • For tcrminolngy anil method* of anntvliiit. *rr «li«rii»^ttiii of ITur o( Itioiiirliiral Aiuil*.;". [•. 71H. SPECIAL TECHNIQUES 101 By the new method the volume of all food items in the specimens examined is added together and then the actual percentage of each is computed. It is as though all the items from all the birds were dumped together and then each kind of food separated and the per- centage of its volume in relation to the whole computed. On this basis, the four beechnuts now equal but 18.18 per cent of the total volume eaten by the two birds. By using this system, a given amount of a single item has the same relationship in all instances. As will be seen by the accompanying table, the differences tend to cancel out where large numbers of examinations are involved. But even with 1,093 grouse analyses, the differences with some food species, figured by both methods were substantial. On the basis of use, four of the species classed among the first 1.5 by the old method were replaced under the new method of computation. Lastly, the "volume" method is quite as easy as the old one. TABLE 98. COMPARISON OF THE "AVERAGING PERCENTAGES" AND "AVERAGING VOLUME" METHODS OF SUMMARIZING GROUSE FOODS Fowls Aspen Cherry Birch Hdspberry Hui>-hurnbeani Thortmpple .Struwberry \pple Beech .Sumach "Based on 1.093 birds. .^Based un :I0R birds. All seasons* Averaffe by volumes Per cent 12.4 10.6 9.2 8.8 .1.7 4.S .3.8 .1.8 .1.0 2.S Average by percentages Per cent 10.3 10.4 7.0 8.8 4.2 6.1 3.8 4i5 3.3 FallA Average by volumes Per cent 6.6 7.4 1.2 6.0 11.8 3.5 6.9 8.8 3.3 Average by percentages Per cent 1.8 6.4 3.3 2.3 3.9 15.9 1.7 4.6 10.3 3.1 It was not always practical, however, to take advantage of the greater accuracy permitted by the use of volumes. For example, the amount of food eaten by a 3-week-old chick is many times greater than the capacity of a chick less than a week in age. Obviously, one must expect an increase in capacity as the bird matures. Since it is impractical to collect, each day, sufficient specimens representative of each successive stage of growth, it becomes necessary to consider larger groupings witli the less desirable, but none the less unavoidable, difference in capacity. For this reason the discussion of the juvenile diet is based upon averaged percentages, otherwise the quantity of food eaten the last half of June would so far outweigh that of the first half as to cause the former to lose all significance. Determining Weather Rel.'VTIonships It has been contended in the literature that adverse weather has been related to some periods of grouse scarcity that have occurred. The weather, however, was not described in other than general terms, such as long hard winters or cold wet springs. Tlie task of specifically relat- ing the various weather constituents to ruffed grouse was therefore attempted along two lines. One series of problems was attacked with data from daily records of temperature, wind, humidity and precipitation ( snow and rain ) taken at the Ithaca station of the U. S. Weather Bureau, 15 miles from the study area at Connecticut Hill. Supplemental information was also secured from readings taken from time to time on the area itself. The minor uncon- 708 METHODS Ai\D TECHNIQUES, formities noted hctweeii the two seem largely attributable to a difTereiice in altitude of 1.500 feet. Each time an iniuialure or adult grouse was contacted, the field leaders recorded the weather information for a second series of problems. These observations were in general terms*, tem- perature being noted, for exam|)lp. as warm, nornial or cold. In this case the basis of compari- son was the average conditions for the ])articular time of year involved. Realizing that the influence of weather was the result of the inter-relation of the various constituents, appropriate statistical methods were utilized in evaluating these interactions. The relationship of grouse mortalitv rates to such items as temperature and rain, was determined l)v means of multiple regression". The results indicated, for example, the effect of precipi- tation on grouse mortality, with the effect of temperature also taken into consideration. The weather conditions, as recorded in general terms by the field leaders, were utilized in determining the relation between this factor and the use of various cover types. Such records, when summarized by seasons and anaKzed by chi-square' made it possible to judge whether the use of various types of cover tended to vary according to changes in weather as recognized by the field leader Determining Predator and Buffer Relationships Determination of the importance of predators and buffers as influences affecting grouse abundance was one of the most difficult problems facing the Investigation. In attempting to establish the true relationships, data were accunmlated regarding ]ircdalor abundance, buffer abundance, food habits of predators and predators responsible for grouse found dead and nest destruction. Predator Abundance and Activity Practical methods of estimating actual predator populations were not known and did not become available during the course of the Investigation. Adequate methods of determining trends in activity, however, were devised for the more important ground predators in 1933. These methods centered around the idea of utilizing fluctuations in the number of jiredator tracks as an index of activitv. The idea of activity is emphasized since it a])pears that the fox. for exam])le. produces tracks in relation to its food supply. Similar populations of foxes |)roduced man\ more tracks when food was scarce than when it was plentiful. Thus it is prob- able that a general increase or decrease in tracks, throughout a period of years, may be an indicator of fox numbers, while variations from year to \ear more iicarix represent activity. Predator activitv trends were obtained during tiic winter period when a tracking snow was present. The various tracks were recorded on the daily notes in a prescribed maimer. Since the amount of field work varied between \ears. it was necessary to reduce each year's data to a com(>arable base through the use of a slatisticr scrlion and tiie mimber of times tiie section was adjusted to a base conmion for liolh ty])c and amount of effort. Thus accurate \carlv fluctuations in the abundatice of predator tracks were estab- lished. . -• • S«c fifiiircii T.t til 7(i. A See HiscUMi'iii iif Ki-Kn-iminn jiriil CiirrrUlinn, p. 720. t See dincussion n( Chi-aqtiiri*. p. 719, SPECIAL TECHNIQUES 709 Buffer Abundance Rabbits, mice, and to some extent, squirrels, are the principle buffer species with respect to grouse. When these foods become scarce, predators are forced to turn to other sources and grouse suffer accordingly. Thus fluctuations in the abundance of buffers are of importance in evaluating the effect on grouse of changing numbers or activity of predators. It was found that rabbit, squirrel, mouse and shrew tracks were too numerous to be accurately recorded by the regular survey crews. Accordingly, a controlled tracking system, based on trail counts, was organized. Trails, each one-half mile in length, were laid out through representative cover types on a study area*, care being taken that the locations chosen would change as little as possible in cover relationshi|)s during the course of study. Each trail was marked in such a manner that an identical jtath could be followed during each working and in subsequent years. Tracks were observed and counted along the.se trails in a controlled manner. Approxi- mated 24 hours after a fresh snow, the trails were traversed })v a single man and alwa\s in the same order. In practice, a day was judged to be suitable if it had stojjped snowing the previous day between the hours of 10 a. m. and I p. m. The character of weather during the time since snowing ceased was recorded, together with the character of the snow. All tracks which crossed or touched the trail made by the oi)scrvers" feet were counted as individual tracks. If an animal walked direclK along the trail, it was counted as a single track, but if it deviated from the trail h\ more than one foot and then returned to ihe trail, it was counted a second time. This was done to eliminate the personal element to ihc greatest possible extent. The trails were co\ered as man\ times during the winter period as the above-mentioned snow conditions occurred. Ilowe\er. an attempt was made to cover the set of trails at least four times each year. Trail track count data was analyzed bv the method of covariance by adjusting the number of tracks recorded for the number of times worked. Individual trails were used for repetition. Food Hahils oj Predators In order that the true iniportam e of predators be understood, a picture of the seasonal and yearly changes in their normal food was necessary. This was obtained through the analysis of stomach specimens from the State as a whole as well as drop])ings and j)ellets collected b\ the regular survey personnel on the study areas during all seasons over the entire period of the Investigation. Collections of the latter t\pe were often large, with such items as fox drop|)ings (.scats) sometimes running to 600 specimens for a single season. When examined and sununarized. \earl\ and seasonal clianges in the occurrence of items eaten were revealed. Correlations were also made with known fluctuations in the abundance of some of the more important foods, such as rabbits and mice. This material was examined b\ the laboratory personnel conducting the food habits studies. Predators Responsible jor Grouse Rills and .XrsI Des/ruction^ An accurate knowledge of the predators responsible for the observed mortality of grouse during the stages of their develo|)ment is also of importance. Early in the Investigation, * Eight such trails were used on the Connecticut Hill area. A By R. W. Darrow. 710 METHODS A^D TECHNIQUES preliminary experiments were run at the Research Center to find a method hy which the signs left where birds were killed or nests broken up rould furnish a I)asis for recognizing the predator responsible. Observations of penned wild animals checked by an occasional experience in the field when predators were surprised in the act of breaking up a nest, formed the basis for the following key. One must remember, however, a key of this kind must be confined largely to typical examples while in prartire one frequently enrounters situations where the evidence is conflict- ing. This is particularly true with respect to eggs destroyed toward the end of the incubation period. In such cases, the different indications must be weighed against each other as well as against circumstantial information such as the presence of tracks or hair. Also, the relative probability of the various suspected species finding the nest location must be considered. Beyond this, one may record the quality of the evidence by rating the diagnosis as excellent, good or fair. Furthermore, the key deals only with the characteristics of wild predators, except that the dog and house cat are included, and also applies only to species found in New York. It assumes that nests destroyed as a result of such causes as fire, flood or the activities of man will be recognized by the investigator from other sources. Key for Diagnosing Predators Responsible for Breaking up Grouse Nests A. All or most of clutch intact in nest 1. All eggs in nest, cold Female probably killed* 2. Three or four eggs missing; remainder usually still being incubated Foxes frequently filch a few eggs at a time from a nest. Crows and red squir- rels may do the same thing. Further- more, any species which carries the eggs away from the nest may be interrupted and thus leave this result^ B. Nest empty of eggs or shell remains but structure not disturbed^ 1. Remains of individual eggs left at points from 1 to 1?> feet from nest in different directions a. Shell usually retaining its general spherical form n) Shell irregularly opened at one side: frequently found on stumps (a) Punctures usually conforming to shape of crow's bill, frequently with smaller out- ward punctures opposite Crow (b) Opening not having punctured ajipear- ance; two portions of shell sometimes separated : yolk or other egg contents occasionally found smeared or tree trunk: all eggs seldom taken Red Squirrel * If the rrnminii ran lir (oiiikI iIii'v may give Boine indication of thr |>rr<]alnr respnnftiblc. A Soc B for rharnrlrriKtir melltndii of tlip«c ipteivf. t In flfldition to Ihr Rprrieii limn! nndrr llti* rstrsory the blnrk snnkr, wliirli occniiionally Iihb lircn fount! guilty. vrouM Iravi, the nml In Ihia condition, Hrrauiir it awallowa the CRpt whotr. ratrliinp tllP individual Willi ila loot it the only way of apprrlipnd- in|[ it. SPECIAL TECHNIQUES 711 (2) Shell having a large "bite" taken from one side about midway between the ends and extending from one-half to three-quarters the way around; opening sometimes smaller but inward punctures made by the canine teeth can usually be found in the opposite side Red Fox, Gray Fox b. Shell not retaining shape, one-half or more usually eaten away along the long axis; edge of remaining portion very finely chewed but the fragments large- ly held together by the membrane; shell remains frequently left under a log, projecting root or sim- ilar shelter, often several at the same place Weasel*, Mink 2. Whole eggs buried in runways under or near nest Mice^ C. Shell remains in or close beside nest, latter often torn apart 1. Nest torn apart, completely crushed eggshells mixed with the debris Skunk 2. Nest intact or very slightly disturbed a. Shell remains usually retaining much of their spherical shape, one end bitten oS, most frequently the smaller one; if broken to a greater extent the fragments neither held together by membrane nor mixed with debris Raccoon 1). Shell remains well crushed (1) Remains, much chewed but fragments held to- gether by nieiiibrane, largely in nest, latter not disturbed Bobcat, House Cat (2) Remains largely absent, but small bits of shell usually scattered in and beside nest Dog, Opossum, Woodchuck^ * Either Musu-la novaboracensis or M. cicognanii, A Except that they may have been involved in some of the few instances of this kind, the Investigation has found no evidence that chipmunks disturb grouse nests. t Little dilTercntiatioii is possible between these species except on a circumstantial basis. Dogs mure frequently paw up the nest but lliere is considerable variation between breeds as well as individuals. The majority of broken-up nests were examined only by the authors throughout the In- vestigation in order to eliminate the personal element to the greatest extent. Their reports, together with all the evidence that could be gathered was in turn checked by one individual to reduce further the possibility of error. The same author also decided which predator, if any, was responsible for each grouse found dead. Tlie field evidence was noted on an appropriate data sheet. The feathers, bones or other remains were placed in a bag. The specimen and the sheet were then checked in light of the results of experiments as well as field experience. Unfortunately, judging the predators or cause responsible for adult grouse kills does not lend itself to being summarized in key form since circumstantial evidence is involved, and space does not permit the detailed discussion which would otherwise be necessary. a SPECIAL TECHNIQUES 713 REMAINS OF ADULT GROUSE KILLED BY AN ACCEPITRINE HAWK (PROBABLY COOP- ER's) SHOWING HEAD STILL ATTAGHED TO CARCASS, FEMUR PICKED CLEAN BUT not fractured. and gizzard uneaten Determining Effect of Hunting The proportion of birds taken by the gun and its efFeit on prouse populations has always been a debatable subject. To collect evidence on this a hunting check was established and used in 1930, 1931, and again in 1936. On the first trial the areas under observation were posted with signs asking the hunters' cooperation and giving instructions. Shellacked envelopes con- taining printed postal cards were nailed beneath the posters. The hunters were asked to fill in and return the questionnaires to the Investigation. The following year, instead of ])ostal cards, blank spaces for names and addresses were left on the posters. Personal visits to obtain information were then made to those who cooper- ated. Both of the above methods were dropped when it became apparent that children or un- thinking sportsmen did not cooperate but destroyed or defaced the envelopes or posters. In 1936. with the aid of CCC labor, a method of checking hunters in and out of an area EXPLANATION OF PLATE ON OPPOSITE PAGE. a. Red fox — showing < hiinuterislic nwnrier oi biting into egg from side bnt not crushing it greatly. b. Rarcoon — charaeterislirally bites into end of eg^ (upper row) hut sometimes quite similar to fox (lotver row). c. Skunk — showing high degree to which shells are crushed. d. Weasel — showing typical fine chewing of edge oj shell along longitudinal axis. e. Mink — similar to weasel but somewhat coarser. f. Crow — showing "pecked" nature of holes in shell. g. Crow — showing outward pipping on opposite side. 714 METHODS AND TECHNIQUES by establishing check stations was used. A form sheet was filled out for each man and any grouse taken were examined. The questions asked by each method were identical. They concerned the number of grouse flushed, number shot at, number killed, weight and sex of those taken, number of hunters in the party, number of dogs, if any, number of hours hunted, other game taken, and the probable number of birds crippled. Determining Disease Relationships The birds collected* were utilized for both food habits studies and pathological examin- ation. Most specimens were shipped to the Research Center by express, refrigerated by one of two methods during the warmer months. Birds taken during June, July and August were placed in wax paper bags immediately on collection to retain whatever external parasites might be present. These specimens were shipped with dry ice, when it was available, or in wooden boxes containing wet ice and sawdust. In some cases, blood smears were made by the collectors in the field. When the collector was within driving distance of the Research Center the smears were made at the end of the day of collecting. Those of birds collected by persons not in a position to make them or to bring the specimens to the laboratory, were made on arrival. They were not taken from all birds. The distribution of birds from which blood smears were made is discussed in the chap- ter on disease. Many other grouse, supposedly diseased, and some viscera were sent in by sportsmen and game protectors. Examination of the birds involved the systematic search of all organs for parasites and pathological conditions. Standard methods of collection and preservation of parasites were used. A search for coccidiosis was made by the smear method rather than by the flotation of oocysts. This also revealed the presence of microscopic tapeworms. Identification of parasites was checked from time to time by workers at the U. S. Bureau of Animal Industry. Routine procedure included recording the weight, sex and age of all birds. The crops and gizzard contents were weighed and preserved for food habits examination. Estimating Grouse Populations The determination of the densities of grouse per unit of area was a necessary part of the solution of many problems. As mentioned previously, a complete coverage of each study area was necessary not only for grouse population estimates but also for other purposes. A method for determining their number was, therefore, evolved from the data gathered by this system. It became evident that a single working of an area did not yield contacts with all of the birds present. Previous information, plus data gathered during the first year of the Inves- tigation, however, indicated a characteristic of the bird which aided in the development of an estimation system. Adult prouse. particularly during the spring period, exhibit territorial tendencies. It was found that by working the area three or preferably four times, over a short period, the territories of individuals or groups of individuals could be j)lotted with fair ac- curacy and that the chances of completely missing a bird during all of the workings was slight. The population of a compartment was, therefore, a summation of its occupied territories and the population of a study area was the sum of the compartments. * See difcuMion of Determining Food Kelatiunshipi, p. 70MTH BAKE (.litU Ml iiKNKATII AM) (lower) wire cage TRAP WITH CM'TIRED RIRD SPECIAL TECHNIQUES JV, The method attempted during the winter operated through the attraction of birds by the use of bait. During the late fall, small shelters of evergreen boughs were placed in areas known to be frequented by grouse. These were baited with apples and grain. A specially- designed wire trap was placed in the shelter and. when it became evident that the grouse were eating the bait, the tra|) was set. Each day the snow was swept clean from the ground under the shelter. The success of this system varied considerably. Once a bird learned of the existence of the food it was almost certain to be caught. But if the bird had changed to its winter diet of buds, it would walk through a shelter with the bait in plain view and apparently not rec- ognize it as food. Only a few birds were caught by this method and, after two years, it was abandoned. During the nesting season a method of trapping the female grouse on her nest was de- vised which met with greater success. Once located, the Investigation was able to catch and mark the female in better than 90 per cent of the cases. The trap consisted of a rectangular piece of chicken wire held in an inverted V shape bv a stiff piece of wire at one end. A door was hinged to the top of the L . The trap was set over the nest and staked or nailed down around the back and sides. The door was propped open with a short stick to which a piece of string from 7.5 to 100 feet long was tied. After setting, the trap was revisited in about two hours and the door sprung by pulling the string. The bird which, with very few exceptions, would have returned to her nest inside the trap was easily caught. The flexible chicken wire was pushed down around the struggling bird and held by one person until another could extricate her from the wire. Broken wings seemed to be the major risk but such accidents were very infrequent. A small number of the birds, less than 10 per cent, deserted their nests after being captured. GROUSE TRAPPED AT NEST AND MARKED WITH COLORED CHICKEN FEATHERS WIRED TO ITS TAIL After the bird was caught, marking for field identification was accomplished by ... im ^fCs colored feathers to the tail in various patterns. White, yellow, orange and pink were found *" ■ t i'"^^' ■18 METHODS A \I) TECIINIQVES to be the iiiosl easily identified colors. White leghorn chickt-ii feathers were used. Coh>rs were obtained by using the following commercial dyes: Yellow — Wool Yellow — Extra Concentrated Orange — Wool Orange — 2 G Crystals Pink — Croceine Scarlet M 0 0. The marking feathers were easily visible and usually remained until the tail feathers were moulted during late August. All birds were weighed and aluminum leg bands put on before release. Blood smears were taken, in a number of cases, from a vein in the wing. Determining Sex and Ace Insofar as laboratory techniques were possible, sex was determined by dissection and exam- ination of the internal organs*. External appearance and behavior have also been used as discussed elsewhere'^. No means of determining age beyond one year is known. In order to distinguish birds of the year from adults of more than one year the presence or absence of the bursa Fabricii afforded the best criterion. This is a sacular structure which opens into the cloaca. It is pres- REAR PORTION OF BODY CAVITY OF A BIRD OF THE YEAR (LEFT) SHOWING BURSA AS COMPARED WITH THAT OF AN OLDER BIRD ent in young birds but is absent in adults. Its use as an index of age in game birds was pointed out by Gower"^. Experience has indicated that in grouse the bursa may be relied upon to separate birds of the year from adults through February and that a certain number of birds may show bursae during March and April, though its absence during these months does not necessarily mean that a specimen is not a bird of the year. Another feature which has also been used is the character of the first two primaries, since the juvenile feathers are not moulted until the late winter of a bird's first year'f. USE OF RIOMETRICAL ANALYSIS Throughout this book references have been made to the use of biometrical methods in the analysis of data. When the Investigation was organized in 1930 advanced statistical techniques * Sec Anatoui), p. "40. A Sec Chapter U, p. 39. t See Chaplcr \\, p. 84. USE OF BIOMETRICAL ANALYSIS 719 were not in general use in wildlife research. As the work progressed every effort was made to subject each problem to the most thorough and critical analysis possible. At the same time methods were being developed wherebv the significance of biological data could be judged more precisely. As applicable techniques of this kind became available the Investigation has used them. The results have added weight to the conclusions drawn. Substantiation has been given to many relationships already recognized while new ones have been disclosed and some, appar- ently valid by inspection, but found not to be statistically significant, have been discarded. The wildlife biologist, of necessity, must often work with samples of unknown quantities, whether they be grouse populations, cover composition, food availability or weather. A fun- damental characteristic of such samples is variability, i. e. repeated measurements of the same thing seldom yield the same value. The greater the variability, the greater the number of measurements that must be taken before a summation of the information will not be unduly affected by an additional measurement that might, by chance, be either very low or very high. Statistical methods have afforded a means of taking this aspect of the data into account in analysing the records of the Investigation. Of paramount importance has been the fact that biomelrical analysis provides objective criteria for evaluating the significance of relationships in terms of probability. In other words, they make it possible to determine the degree to which a correlation may have occurred by chance rather than from cause and effect. Significance To denote the degree to which chance may be involved three terms are used, namely, not significant, significant, and highlv significant. These terms define three limits of probability. Not significant indicates that the relationshi|i would be expected to occur more frequently than once out of 20 times by chance alone: significant that it would occur not more than once out of 20 times by chance; highlv significant that it would occur by chance not more than once out of 100 times. The mathematical basis of statistics is not important li> the average field worker, but may be found in standard text books'"' "*• "°. However, a brief discussion of the methods used in analysing the data of the Investigation seems worthwhile. Chi-Square The chi-square test is designed to compare observed with expected or hypothetical values and to determine the significance of departures therefrom. With sex ratios*, for example, an expected ratio of 50 males to 50 females was assumed and the magnitude and consistency of the recorded deviations from this ratio controlled the degree to which such differences might be considered real. Chi-square was also used in evaluating cover type use. Here, however, the determination of the "expected" values from which departures could be measured constituted a preliminary problem'^. Analysis of Variance When data are taken in such a manner that the variability of a single factor can be segre- gated into component ])arts. each traceable to an independent source, then the technique known as analysis of variance may be used to test the significance of each. In preparing this re- * See Chapter VHI. A See discussion of Dclcrniining Sliellfi Relatioii^liips. p. Till, 120 METHODS i\D TFCHNIQUES port it was especially useful in untangling the effects of years, seasons, sex and age. For ex- ample, a series of weights of grouse collected over several years from all seasons would mean little until analysed by this method. When the variability due to each of these influences was separated from that of the whole it became possible to judge the significance of each. Regression and Correlation In many wildlife research problems knowledge of the relationshij) or association between two or more factors is important. Regression and correlation are methods of analysis which provide means of expressing the degree of relationship between such factors. They summar- ize, in mathematical terms, information usually presented by graphs and visually appraised. Mathematical evaluation has the advantage of leading to tests of significance. One of the most important uses made of regression analysis by the Investigation was in the statistical adjustment of data. For example, variations in the degree of overwinter mortality recorded among grouse on the study areas over a series of years required such adjustment with respect to fluctuatiotis in the preceding fall population level to obtain comparable yearly trends. Similarly, records of the number of fox tracks observed were corrected for differ- ences in the amount of effort spent in the field from year to year. Simple regression, when extended bv measuring more than two factors coincidentally. gives rise to a form of analysis termed "nuiUiple regression". When several groups of such data are handled together the analysis of variance is combined with either simple or multiple re- gression in what is known as ''analysis of covariance". This method was. for instance, used to correlate records of wind. rain, man-hours and grouse flushes on the xarious compartments of the Connecticut Hill area for a series of years*. * See diwcussiiiii of Fstiinuting Cnnisf Population!), p. 714. THE ANATOMY OF THE RUFFED GROUSE* By David E. Davts The ruffed grouse is a member of the family Tetraonidae and. like other Galliformes, has a generalized structure, as shown by the presence of two carotid arteries and a gall bladder, and the absence of a penis. The intestines are looped in a primitive manner within the body cavity. There is no hemipterygoid bone. The palate is schizognathous. However, the para- sphenoid rostrum (basjsphenoid rostrum) has specialized knobs for the articulation of the pterygoids which are not homologous with the basipterygoid processes. A notable specialization is a cutaneous muscle lying beneath each tract of ruff feathers. This muscle is about two centimeters long by one centimeter wide at the base and is somewhat triangular with the apex pointed ventro-anteriorly. The feather follicle extends into the muscle, and the contraction causes the rachis to rise away from the neck. Edgew'orth'" states that M. cucullaris is not developed in birds and its functions are taken over by M. cranio- cervicalis. The ruff muscles are probably a derivative of M. cranio-cervicalis. The Skeleton Skull. The skull is illustrated in figure 77 a-c. The premaxillary bones are concave down- ward and pointed anteriorly to form the top of the beak while posteriorly they terminate in three prongs. The nasal processes are dorsal and pass between the nasal bones as far back as the frontals. The maxillary process on each side passes backward along the border of the jaw to form part of the jugal arch. The palatine jirocesses extend on the surface of the palate to join the palatine bones on each side. The maxillary bones lie posterior to the premaxillary bones and each has two processes. The ])alaliiie process lies along the palatine bone and the jugal process forms most of the jugal arch. The nasal bones are broad and are sutured to the frontals. Each has two anterior pro- cesses. The premaxillary process is a thin pointed sliver which lies along the external side of the nasal process of the premaxilla. there being an open slit between the two. The maxil- lary process extends downward to join the maxillary bone. The external nares are holorhinal, formed by the premaxillaries in front and the nasals behind. The nasals are sutured to the frontals in such a manner that a forward extension of the frontal lies under the nasals but is not fused with them. The frontal, comprising the surface of the skull in the interorbital region, is concave up- ward and somewhat narrowed. Behind this lies the parietal, a thick spongy bone which makes up the hinder part of the top surface and extends down to the supraoccipital bone to form most of the posterior aspect of the skull. The latter lies directly above the foramen magnum. These bones of the top and back of the skull are paired during early development but in the mature bird are so completelv ossified that neither the sutures between the |)arts nor between * The sttuly upon which this account is based was undertaken by Dr. Davis in collaboration with the New York State Con- servation Department and under the auspices of the Museum of Comparative Zoology. CambridRe. Mass. The author is indebted to the late Dr. Glover M. Allen for innumerable suggestions and constant advice, as well as to Miss Katherine Wilson who assisted in the preparation of the drawings. .\cknowledgment is also due to the late John C. Phillips for specimens utilized in dissection. 722 THE ANATOMY OF THE RUFFED GROUSE the difFerent bones can be identified. The posterior aspect of the skull is sub-circular. The foramen magnum, bounded by the occipital bones, is round except on the ventral border where the occipital condyle protrudes. On each side of the foramen are two small foramina for the hypoglossal nerve. The glosso- phyaryngeal, vagus and accessory nerves pass out through a more lateral foramen on each side. At the lower lateral margin of the posterior aspect lies the otic region formed by the coalescence of the three otic bones. Just at the edge of this area is a fossa into which passes the internal carotid artery and from which pass out the vena capita lateralis and the palatine branch of the facial nerve. TTie orbit occupies most of the lateral aspect of the skull. Anteriorly, it is bounded by the prefrontal (lacrimal) which is a scale-like bone extending from the frontal — to which, however, it is not ankylosed — part way to the jugal arch. The interorbital septum has a small centrally placed fenestra. The posterior wall of the orbit is formed largely by the or- bitosphenoid and the pleurosphenoid ("alisphenoid"), and is bounded by the postorbital pro- cess of the frontal. This process is fused distally with the zygomatic process of the squamosal bone to produce a sharply pointed forward extension which is serrate on its ventral border. The temporal muscles pass through the canal formed by the partial fusing of these two pro- cesses. The squamosal, comprising much of the lateral wall of the brain case as well as a portion of the orbit, is fused anteriorly to the pleurosphenoid, dorsally to the frontal, dorso- posteriorly to the parietals, and posteriorly to the exoccipitals. There is an otic process of the squamosal which protrudes over the otic region. The olfactory tract lies along the underside of the roof of the orbit and has a large opening into the brain case. The optic foramen, through which passes the optic nerve, is large. The oculomotor, trochlear, the pro- fundus branch of the trigeminal, and the abducens ner\'es pass out through separate foramina. In the pleurosphenoid and below the zygomatic process is the foramen ovale for the maxillary and mandibular branches of the trigeminal nerve. The otic region lies within a deep fossa, its posterior border being formed by the parotic process of the exoccipital and its anterior border bv the pleurosphenoid. Tlie anterior part of the fossa is a reticulum through which the Eustachian tube passes downward from the in- ternal ear to the mouth. Tti the sidewall of this tympanic cavity is the fenestra ovale, an obvious opening in which lie^ the columella. Tlie facial nerve docs not come out on to the surface of the skull but. with the vena capita lateralis. j)asses through a bony canal just supe- rior to the fenestra ovale. This canal opens to the exterior just medial to the position of the internal carotid artery in the fossa on the posterior aspect of the skull. The semi-circular canals arc not imbedded in solid bone but lie in a spongework. Air enters this area through the Eustachian tube and reaches the parietals and other skull bones through pneumatic openings. The j)alate is perfectly schizognathous. The pterygoid extends from the quadrate to the rostrum of the parasphenoid. where it articulates with the palatine. This latter bone, on each side, extends anteriorly, wide at first, and then narrowing to join the palatal processes of the premaxilla, while its posterior external angle is completely rounded off. Tlie maxillo- palatines pass dorsad of the jialatine and are directed ])osteriorly and medially. The rostrum of the parasphenoid is thick and strong. The i)revomcr ("vomer") consists of a pair of carti- lagenous slivers just anterior to the rostrum. The basijitcrygoid processes are represented by two facets on the paras])henoid rostrum. Tli<' paras|)heiioid has two ])airs of openings in its anterior part. The medial pair is for the Eustachian tubes and the more lateral ones for tin- internal carotid arteries. Tlic basis crnnii is roniuh'd and bulcring. i Q. So J ^g; 60 -J U. ^ H C 0, = -,•<=> a. o -B fe 5 I <;=>. 724 THE ANATOMY OF THE RUFFED GROUSE The quadrate bone consists of three processes. The otic process has one condyle, as in most ])iimitive birds, and articulates with the squamosal, thus forming the main articulation of the jaw. The orbital process protrudes into the posterior region of the orbit. The ventral process is divided into two parts, one of which articulates with the lower jaw and the quad- rato-jugal arch, while the other is more medial and articulates with the pterygoid. The external aspect of the lower jaw is formed from the dentar\ . angular and surangular. The dentary forms the most anterior part and extends posteriorly to the fenestra between the surangular and the angular, and on the lower edge passes beyond the fenestra for some dis- tance. The surangular meets the quadrate. From it there are two processes: the internal ar- ticular process extends into the orbit for a little distance: the retroarticular process curves upward around the posterior part of the quadrate. The latter is a long process on which is inserted M. depressor mandibulae. On the internal aspect of the jaw, the splenial bone is discernible, covering the inner side as far posteriorly as the fenestra. The other elements of the lower jaw are completely fused to the angular and surangular. In general, the skull of Bnnasa is much lighter than tliat of nther members nf the Tetraoni- dae. The skull of Lagopiis is proportionallv shorter, broader and higher, andr' has no slit between the nasals and the nasal process of the premaxilla. Centrocercus has a highly arched bill and low frontal region. The nasals are as in Lagopus. The prevomer ossifies, although it is verv small. The mandibular fenestra is slit-like. Tyinpaiiuchiis has a large prefrontal bone, a long postorbital process and the prevomer is ossified. Vertebral Column. There are fifteen separate cervical vertebrae, including the atlas and the axis. The atlas, which bears the head liv means of a socket into which fits the occipital condyle, is a ring-shaped bone possessing poslzygapophvses and an articulation on its lower posterior surface for the odontoid process of the axis, upon which the whole revoKes. The postzygapophyses of the axis form a wide flange of bone. The next two vertebrae have this flange of bone ])enetrated by a fenestra, as in Galliis. The following six vertebrae are elon- gated and have the renniant of a rib fused to the diapophysis and directed posteriorly. Their postzygapophyses are long and thin, bearing their articular facets at tiic tip. The next {\\f vertebrae are shorter and stouter. The second I axis I. third and fourth vertebrae each pos- sess a sharp, keel-like hv|)a|>o])h\sis. On the fifth, the li\ pa|)ii|)h\sis is a small narrow process. In these characters Boiiasa differs from Galliis in «lii(li tlii> hypapophyses are long and sharp"'". The last four cervicals each have an anteriorh direiled hvpa|)o])hvsis. but the sixth to eleventh vertebrae imhisive lack hypapophyses. The terminal ccr\ ical \ertelira possesses a comparativeI\ short, thin, cervical lib which is articulated willi imili ilic rcniiurn and the dia- ])ophvsis but which does not reach the sternum. The traiisM-rse foianieii is progressively larger till, in the last cervical vertebra, it is lacking. There are five thoracic vcrteiirac of wiiirli the first four arc fll^ed. DorsalK. there is a ridge formed from their fused neural spines. The tips of the first three hypapophyses are fused, thus forming two fenestrac on the ventral side. The tip of the middle hyjiapophysis is ex|)anded to form a small plate, as in Tympanuchus. In Callus, the first fenestra is smaller. The tips of the transverse processes are idimcclcd and the foramina thus formed are partK filled by a thin film of bone. The articulation for the head of the rib is clearly visible on the last three thoracic vertebrae. The free llioraiil seapulare externum arises on tiie external fare. This last head is small. M. triceps is the great extensor of the ulna and radius. It is divided into three heads. Caput THE ANATOMY OF THE RUFFED GROUSE 737 scapulare is the largest and originates from the internal basal part of the humerus and the adjoining faciae, and passes fleshy along the humerus between the M. deltoides major and the body. Furbringer"" classifies the origin of this head into four types. Bonasa belongs to type 3 — "proximale humerale Ankerung". Caput coracoideura consists of a long tendon which originates by a branched tendon from the M. coracobrachialis posterior and M. proscapulo- humeralis. and passes along the ventral surface of the wing to end in a small muscle which expands the secondaries. Caput humerale originates on the dorso-medial surface of the humerus and passes along the medial surface to insert on the olecranon of the ulna. This head is attached to the humerus for its entire length. Specialized Structures and Systems Syrinx. The syrinx is formed by modification of the junction of the trachea and bronchi, and thus is of the tracheo-bronchial type. The last six tracheal rings are reduced in size but are not vestigial and are complete rings. In the sub-order, Galli, each tracheal ring is fused on the dorsal side and thus does not leave a drum-like area^'". The penultimate ring is somewhat enlarged ventrally to form a small ixi-lcriorlv directed protrusion and dorsallv it is fused with the dorsal plate of the pessulus. The ultimate tracheal ring is fused with the ventral plate of the pessulus but is incomplete dorsally. The first bronchial semi-ring is not attached to the plate of the pessulus but is suspended by the mcmbrana tympaniformis externa. Its dorsal end is free and expanded. The second bronchial ring is attached to the ventral plate of the pessulus and is wider than the first ring. The pessulus is rod-like in form and extends dorso- anteriorly. Its ends are expanded to form a dorsal and a ventral plate. The membrana tvmpaniformis externa forms the lateral wall of the syrinx. The mem- brana tvmpanifcji mis interna strctchi's behveen the ends of the bronchial semi-rings. The bronchidesmus is a membrane stretched between the membrana internae of the bronchi. Triangular pieces of cartilage are included within the membrane. There are two sets of muscles which move the trachea and syrinx. The M. trachealis set is composed of one muscle, extending along the trachea as far posteriorly as the sixth (last) tracheal ring. Its function is contraction of the trachea. The M. sterno-trachealis set is composed of one muscle on each side of the syrinx. The insertion of each member extends from the penultimate trachael ring anteriorly along the dorso-lateral border of the trachea for about two centimeters. A slip extends posteriorly to cover the dorsal plate of the pessulus. The origin is on the outer lateral process of the sternum. Digestive Tract. This is illustrated in figure 80. The mouth cavit) contains the thick muscular tongue, which is broad, short and triangular in shape. On the floor and roof of the mouth are numerous short, hardened sjiines pointing posteriorly. There is a definite row of these surrounding the entrance to the pharynx. The oesophagus is a simple tube extending to the crop. Although Tuttle*^ mentions rudimentary vocal sacs, there is no indi- cation of an extension of the oesophagus related to the production of vocal sound as in Tympanuchus™. The crop is a large spherical diverticulum of the oesophagus lying between the divisions of the furculum on the upper part <>f the breast, mainly on the left side. Its opening to the oesophagus has appro\imatelv the same diameter as the oesophagus itself. Gallinaceous birds have a true crop, which is a diverticulum of the oesophagus, rather than a "false crop" formed by the expansion of the oesophagus as found in certain hawks. From the crop, the oesophagus extends as a short tube to the ventriculus. 738 THE ANATOMY OF THE RUFFED GROUSE ~ ;j(f,t-_ Trachea *r-Z a. I. Mouth 2a. Esophagus ■1 b 2b. Crop *-a c 2c. Esophagus 3. Proventriculus 4. Gizzard 5. Liver 6. Duodenal loop 7. Spleen 8. Pancreas 9. Small intestine 10. Caeca I I. Caecal junction with large intestine (rectum) 12. Large intestine (rectum) 13. Cloaca 14. Vent ir.V/i.im ;/. I.nii - FIGURE !>(). DIGESTIVE SYSTEM The latter is divided into an anterior glandular portion (proventriculus) and a posterior grinding jjorlion (gizzard), with a narrow i.-^tlinuis separating tbein. Gallinaceous birds lielon^ to the tyjie in which the jiroventriculus is separated from the gizzard by a section which is non-glandular internally""'. These birds have the ventriculus highly s])eeialized, for feeding on vegetal material, by the great developincnl of llic niDsciilar |)(>rtiiiii and the tough thick inner lining. The duodenum extends fidiii the (((■r>;d and anterior |)ail of the gizzard and then forms a longitudinal loop. The pancreas lies between the arms of this loop. The entrance of the |)ancreatic duels marks the beginning of the small intestine. This passes immediately dorsad THE ANATOMY OF THE RUFFED GROUSE 739 and then forms several loops, the last one extending ventrad and at right angles to the axis of the body. It then passes dorsad and anteriorly to join the large intestine. The two caeca diverge at this point and form a series of coils at right angles to the body axis, posterior of the gizzard. The small and large intestines are of equal diameter. There is considerable varia- tion in the details of the arrangement of these structures. The Galli are grouped among the birds with long caeca as opposed to the types in which they are small or functionless^". The large intestine (figure 81) is the shortest section of the gut. It lies just under the roof of the visceral cavity and is divided into three parts, coprodeum, urodeum and proctodeum^. C/JRCTAe EPioEQtnia PaocTooLOAt FIGURE 81. LONGITIDINAL SECTION OF LARGE INTESTINE Ix 0.21 I The coprodeum extends from the caeca to the small flap which separates it fnmi tlx- urodeum. The latter is a short section, bounded posteriorly by a flap. It receives the ureters as well as the oviduct or the vasa deferentia. The proctodeum is the terminal section. On its dorsal side is a diverticulum, the bursa Fabricii*. The external opening is smaller than the cavity of the proctodeum. TABU'; 100. AVKRAGE MEASUREMENTS OF DIGESTIVE TRACT OF U. XDILT (JROUSE Part Average Meusurcmeat Oesophagus — anlRrior to crop Oesophagus — posterior to crop 1 15 mm. 5:j 1 12 70.1 95 .326 323 Urogenital System. The urogenital system is extremely uniform throughout the Class Aves and is composed of the sex organs and their ducts, and the excretory organs. The kidneys are paired structures lying ventral to the sacrum and dorsal to the aorta and its large branches, the ischiadic arteries. Each kidney is lobed and sends the excretory products to the cloaca through the ureter. * This is a sac-like pouch which is present in birds of the year np to an age of about nine months. It is therefore the most reliable means of separating young from old birds up until alioul March 1. During the latter part of this period it becomes progressively smaller and finally disappears, although it persists later in some individuals than in others. 740 THE ANATOMY OF THE RUFFED GROUSE In the male, the testes are oval bodies lyin<; ventral to the anterior part of the kidneys. In the breeding condition they are much enlarged. The epididymis i.s a mass of fine tubes con- necting the testis with the ductus deferens which enters the cloaca just ventral to the entrance of the ureter. The sperm pass from the testis through the epididymis and ductus deferens to the cloaca. In the female, the sex organs consist of the ovary and the oviduct. Normallv. onlv those on the left side develop. The ovary is in the same position as the testis and resembles a bunch of grapes in appearance. The egg is extruded from the ovary into the infundibulum of the oviduct. The chalaza is produced here, while the egg white is formed in the tube region, and the shell and color are added in the uterus. These regions are poorly defined and cannot be distinguished in the non-breeding condition. Abnormal conditions of the gonads are not infrequent. An ovotestis may develop or, under certain conditions, the ovarv may atrophy on the left and then the right gonad develop into a testis. Such abnormalities often have remarkable effects on the plumage and other secon- dary sex characters. EXPLANATION OF ABBREVIATIONS USED IN FIGURES 78 & 79 Ace Add. pro. Add. sup Amb Biceps Caudofem Cor. brach. post M. Cr. lat M. Cr. med. M. Delt M. Dors, soap M. Ext. d. coram. M. Fi-ni F. prrf. n F. prrf. HI F. prrf. IV F. p. p. II .. M. ..\I. ...M, ...M, .M, F. p. p. ITT M. F. i>r..f. ..„ M, accessorii m. obturatus adductor profundus adductor siiperficialis ambiens biceps candofemoralis coracobrachiali^ posterior flexor cruris lateralis flexor cruris medialis deltoideus dorsalis scapulae extensor digiloriini coui- munis femnrocaudalis flexor perforaliis dipiti II flexor perforalus digili III flexor [>erforatus ilii;ili I\' flexor perforans et perfnra tus dipili II flexor perforans el perfnr:i- tus di^iti III flexor profundus Cast. _ M Cast, ext . ..M. Glut .. M. 11. ant. . M. 11. fern. _.„. .. ..M. 11. fib M. 11. tib. ant M. 11. tib. lat. M. Ischiofem. _ M. Obt M. Pect. M. Per. prof. M. Per. sup M. Pir. .. M. Psoas . .M. Supracor M. Tib. ant. M. Tib. cart .. .Til Triceps -_ M. V. lat. ext M. \'. lat. med. — M. \". med _ M. gastrocnemius gastrocnemius, pars externa gluteus profuncbis iliactis. pars anterior flexor iliofemoralis extensor iliofibularis extensor iliotibialis anterior extensor iliotibialis lateralis flexor ischiofemoralis obturator pectoralis superficialis . peroneus profundus peroneus superficialis piriformis . psoas . supracoracoiileus . tibialis anticiis bial cartilage . triceps . vastus lateralis, pars externa vastus lateralis, pars niedius va-^tus niedius THE PTERYLOGRAPHY OF THE RUFFED GROUSE* By John E. Trainer The pterylography of a bird concerns the arrangement of the feathers on its body. Among the different kinds of birds there is considerable variation in this characteristic, a feature used extensively in studying the relationships of the various groups. Among gallinaceous birds there is great similarity of arrangement and among the grouse alone it is even more uniform although there are minor differences. In almost all kinds of birds the feathers grow in dis- tinct tracts rather than indiscriminatelv over the surface of the body. The intervening areas of bare skin, termed spaces, are covered by feathers situated in these tracts. There follows a description of the various tracts and spaces (figure 82), together with a discussion of the types and numbers of feathers, their variations, and a description of the "snowshoes". The Tracts The tracts in which the feathers are organized are bilaterally symmetrical. Within them the feathers are generally arranged in two series of rows, one at an angle to the other. The pattern is readily seen on a |)luckcd bird although in this study it was found that greater accuracy resulted when the feathers were clipped. Capital. Included in this tract is the entire head region with the exception of the area between the lower jaws. Within it there are a number of small unfeathered spaces which are generally covered by feathers above and in front of them. On the top of the head one series of rows radiates in all directions from the occipital region while the other makes up a set of concentric circles. Both gradually unite with the spinal trad posteriorly. On the side of the head one series is more or less horizontal and the other vertical. In the region of the eye and ear the rows are more numerous and their feathers are highly modified. In the loral region and the forward part of the malar the feathers project nearly perpendicular to the skin, although their tips curve backward. 0\er the rest of the head the feathers generally point backward at their base. The crest originates in the region between the forward pari of the eyes and is made up of about fourteen enlarged feathers. Usually two are especially long. The crest of the female is only slightly smaller than that of the male. Spinal. This covers the upper surface of the bodv fr(iin the capital tract to the caudal. It is divided into the cervical, interscapular, and posterior regions. The same general plan of feather arrangement is present throughout, the major series of rows sloping posteriorlv away from the middle line. The secondary set of rows is not as evident, particularly in the cervical and interscapular regions. In the former, the feathers intermingle with those from the under side of the neck closing off the lateral cervical sjiace. The feathers increase in length posteriiirh . The interscapular region is narrow because of the adjacent space which is apparently needed for freedom of wing movements. The posterior region resembles an inverted wide-bottom vase and encloses a large portion of the spinal space in the middle * The study upon wliich this aci-nutit is l)asfd wns rarried out I>y Mr. Trainer in connection with meeting the require- ments for a Master of Science degree at Cornell Univeijily, 742 THE PTERYLOGRAPHY OF THE RUFFED GROUSE line. The thick, lonp. aftershafted feathers in the interscapular and ])c>sterior regions provide excellent protection. Oil Gland. This tract is small, consisting of eight small feathers arranged in a circle sur- rounding the apex of the oil gland. Caudal. The caudal tract is made u|) of the large rectrices. their coverts, and the feathers surrounding the anus. The rectrices make up the tail proper, although the middle pair has been crowded out of line dorsally. Usually eighteen are found, but deviations from this num- ber are not particularly rare. Birds having twenty are usually large males while those with sixteen are usually females. The number is not associated with age since twenty have been found in young birds. Neither is it a subspecific character. Protecting the bases of the large tail feathers on the upper side are the upper-tail coverts. These lie in a single row and usually comprise eight pairs, occasionally nine. They separate two rows of small, down-like feathers. At the base of the tail, underneath, are the under-tail coverts consisting of about five rows. Those of the first row (nearest body) are very small. The second, third, fourth and fifth rows contain much larger feathers, but gradually decrease in size in that order. A space occupies the middle line. The anus is surrounded bv small tuft-like feathers. Venlral. The entire ventral surface of the body, from the lower jaw to the anal region, is included in this tract. In the majoritv of birds it continues posteriorlv over the breast and CORONAL REGION OCCIPITAL REGION - POST- AURICULAR REGION CERVICAL REGION HUMERAL TRACT SPINAL SPACE FEMORAL TRACT CRURAL TRACT "PEOAL" TRACT INTER-RAMAL REGION SUB-MALAR REGION CERVICAL REGION INFERIOR SPACE DORSAL VIEW FICURK !!2. FKATIIKK TRACKS AND SPACES OF TIIK Rl FFED CROISE THE PTERYLOGRAPHY OF THE RUFFED GROUSE 743 sends a lateral branch toward the wing but in the rufied grouse, as with most other gallinace- ous birds, the main portion seems to extend laterally. This lateral branch is divided into the sternal and axillar regions. The middle portion, also highK developed and well separated from the lateral branch, is the abdominal region. The remaining portions of the tract com- prise the inter-ramal, sub-malar and cervical regions. The inter-ramal region bears feathers which are erect or nearlv so. At the edges they bend outward, mixing with those of the malar region, thus covering the sub-malar space on the lower jaw. The sub-malar region is not well marked off from the inter-ramal and bears similar feathers except for a gradual increase in size. The cervical region which extends to the base of the neck encloses the inferior cervical space on the middle line. The ruff consists of about five rows near the mid])()inl of the neck in which the feathers are more numerous and much larger. They are unique in their silky, truncate tips, often with iridescence. The male differs from the female in the greater number of feathers making up the ruff and usualK in their greater length. The feathers of the sternal and axillar regions protect the larger portions of the ventral surface, primarily because they are much longer than those of the abdominal region. The latter, while well separated from the other two, is poorly defined on the sides where the lateral space of the trunk is more or less obliterated by small feathers. Thus it is, in a sense, continuous with the tract of the leg. In life, however, the small feathers probably have very little effect. In the illustration they are represented by the smaller marks. A gradual in- crease in size posteriorly is well marked. Likewise, and possibly associated with decreased protection from the sternal and axillar regions, there is a great increase in the proportion of downy parts in the feathers of the posterior portion. Humeral. This bears the scapular feathers which cover the adjacent spaces as well as the bases of the feathers of the spinal tract. Alar. Including all the feathers of the wing with the exception of those in the humeral region, this tract is most important from a taxonomic standpoint and is also interesting, due to the fact that in many cases feathers can be recognized as individuals and are quite con- stant among related birds. Roughly, it is made up of the large flight feathers, the coverts and the alula feathers. The flight feathers or remiges consist of the primaries attached to the "hand", the secondaries attached to the forearm and a single carpal remcx between the two. The coverts are regular rows of smaller feathers in a shingle-like arrangement protecting the bases of the remiges as well as of the other coverts. They become progressivelv smaller to- ward the forward part of the wing. The alula might be compared with the human thumb although it actually is the renmant of a second finger. The relative lengths of the remiges produce a short wide wing suitable for the well-known short bursts of rapid flight characteristic of this bird. Among the ten primaries the seventh and eighth are longest and the first is shortest, counting from the innermost. The secondaries are more or less variable in number. Usually there are sixteen or seven- teen, although, because the innermost are relatively small, oidv about fifteen are actually functional as flight feathers. These smaller remiges are not attached to the forearm but in the elbow region and are thus often referred to as tertiaries. Even though the first or outer- most secondary is reduced, a gap in the wing does not occur because it overlaps the adjacent primary. The remaining secondaries are all nearly equal in size. The greater primary coverts, like the primaries, number ten. They lie between the latter, 711. 77/ A riEHlLOGRAPHy Ol THE RIFF ED GROUSE cath distal to the one it covers, the last or outeriiiust being \ery iiiueh reduied. The middle primary coverts are attached near the inner ends of the flight feathers, those of the first two being absent and the remaining ones all small. The lesser primary coverts consist of four or five small feathers associated with the outer remiges. The small size of these middle and lesser coverts is probably linked with the covering of this area by the alula feathers. The greater secondary coverts have their origin proximal to the secondaries, the number of which they usually exceed by one. All have a distal overlap. The first nine are similar in size, the tenth to the twelfth are larger, but from the twelfth on they again decrease. The middle secondary coverts are not distinguishable from the lesser coverts except by their ori- gin. Usually they are two less than the number of greater coverts, with the deficiency at the inner end. The lesser coverts of the upper surface of the wing may be divided into two groups, a set of three rows which functionally belong to the secondary covert series and a number of other rows further forward which simply serve to cover that area of the skin and are included in the term lesser coverts for lack of a better term. The marginal coverts are short, stiff feathers attached to the forward edge of the wing. In the region of the hand their counterparts are tlie carpo-metacarpal coverts. On the under side of the wing none of the coverts are as large as their counterparts on the upper side. The greater primary coverts in this area are about equal in length except for the outer five which gradually decrease. The single middle covert is associated with the first primary. The under lesser coverts are situated in a single curved row. The greater secondary coverts on the under surface lie proximal to the bases of the sec- ondaries and cover their bases. They are longer in the region of the elbow. The under middle secondary coverts are reduced in the ruffed grouse to the extent that they are not exposed but covered by the first row of lesser coverts. Associated with this is the fact that they are not pigmented except at the tips. The lesser coverts, like those of the upper surface, are of two groups. The rear group consists of two rows closely associated with a third and separated from the other by a space. Among the inner lesser coverts are the large axillary feathers. On the under side of the wing an interesting condition exists in the greater and middle coverts of both the primaries and secondaries. These feathers are inverted, with the surface of the feather, normally lying against the skin, here exposed. This is evidenced by the greater amount of pigment on the hidden surface and the presence of the aflersliaft on the ex|)osed surface. Similarly, they curve outward instead of inward. Thus, the morphological dividing line between the upper and lower surface of the wing is between the middle and lesser coverts of the under side. The carpal remex is a greatly reduced (light feather of the wrist which has apparently been crowded out in the bending of the wing. It measures onlv aliout an inch in length and is generally weak. An upper and lower covert, similarly reduced, arc also present. The alula or bastard wing carries four large feathers and a number of coverts. The large series tapers abruptly with the largest outer one reaching to the tips of the greater primary coverts. The alula coverts are continuous with and resemble the marginal coverts and the carpo-metacarpal coxcrts. Ffmoral. This is very well dcvelo|)ed in the ruffed grouse, extending oxer most of the lateral surface of the thigh. Its general shajie is triangular with the apex at the anterior end. From this point the rows radiate posteriorly while several additional rows occur in THE FTERYLOGRAPHY OF THE RUFFED GROUSE 745 the posterior region. Another set of rows curves up toward the back. Besides covering the thigh, the feathers aid in protecting the side of the abdomen by extending backward to the tail coverts. Crural. This tract covers the tibio-tarsus and may be divided into internal and external regions, both of which are similar to and continuous with the femoral tract. The rows are not very definite. The feathers are sparse and present a peculiar appearance, more like fur than feathers. They are possibly of a type which is less subject to wear than the ordinary flat, stiff kind. The feathers of the tarso-metatarsus, although a direct continuation of those of the femoral region, are sometimes considered to make up a separate pedal tract. Thev are most prominent in the more northern subspecies*. The Spaces As already mentioned, around and between the feather tracts are areas of bare skin called spaces. Although devoid of feather growth'^ they are, in life, covered by the feathers adja- cent to them. Like the tracts, their arrangement is similar on both sides of the body. The following have been recognized. Capital. Several areas of this kind are found on the head. The large temporal space is above and in front of the ear. It is covered by feathers on either side and in front. Above the eye is the superciliary space into which folds the upper eyelid. A similar but smaller space is found on the lower eyelid. Ordinarily these two spaces arc not covered by feathers. The rictal space extends over the rear part of the lower jaw ])ciiic on either side. In the middle of the forehead is the frontal space. Lateral Cervical. This space begins about inidwa) in the fr of the trunk. The functions of the spaces have been described above when quite deliniteU accepted. Some, however, are without any definitely known function. For several of these, possible explanations present themselves. For example, certain spaces of the head and the inferior space occur on protruding bones. It is generally conceded that these spaces provide a smooth contour but possibly the bones pressing against the skin hinder an adequate blood supply. Again it is probable that a number, such as the spinal space, facilitate increased efficiency through allowing greater centralization of the blood vessels supplying the feather tracts. The Brood Spot. This might also be considered as a space. In order to incubate the usual clutch of eggs, practically the whole under surface of the body is plucked of feathers. All feathers, regardless of size, are removed except, apparently, the hair feathers. The area plucked roughly includes the last four rows of the sternal and axillar regions and extends from there almost to the anus. Often the forward edge is irregular. On the legs, all of the area which could come in contact with the eggs is plucked. The Feathers The feathers of any bird comprise a variety of types modified for divers purposes and differing widely between species. Those found in the ruffed grouse are described, as well as what little is known concerning total number and variations with sex and season. Types. The feathers which make up the downy coat of the newly hatched chick are re- ferred to as neossoptiles, being characterized by the presence of pigment and the absence of a shaft. They range in length from 2 to 14 mm. With certain exceptions, the neossoptiles are followed by the juvenile plumage, a fact which has led to the application of the term me- soptiles to the latter. In addition to their manner of development, a weaker, lacier appear- ance and pale coloration also distinguish the mesoptiles from the typical feathers or teleop- tiles of the adult. Filoplumes are the so-called hairs, actually highly modified feathers. They are distributed generally over the body and can be readily identified growing from the elevations of the skin surrounding the bases of the ordinary feathers. Plumulae, or true down feathers, are characterized by the absence of both a central shaft and i)igment. They are. strictly speaking, entirely lacking in the ruffed grouse although a number of small semiplumes have the superficial appearance of plumulae. Such feathers are found in the various spaces and among the coverts of the wings and tail. The tyjjical feathers are the teleojjtiles coni|)rising the rectrices. remiges. their coverts, and the general contour feathers. Basically, each consists of an inner, fluffy portion for warmth and an outer flat, stiff portion, but variations range from the almost completely pennaceous rectrices and remiges to the almost completely downy feathers found anterior to the anus. Many of the highly modified leleoptiles are functional although some are probably only vestigial. Those of the evelids serve as small eyelashes. The feathers of the tuft on the oil gland are without a central shaft and are greatly weakened. The auriculars covering the ear openings are specially adapted for the purpose of protecting the ear without interfering with hearing to any extent, the stiff shaft lu-aring very few barbs. Com|)leteK (li>«n\ Icicdptiies are (]nile numerous and are frequentl\ reft i red to as semi- plumes. Unlike tile true down feathers, the) possess a central shaft. Seldom are they exposed THE PTERYLOGRAPHY OF THE RUFFED GROUSE 747 to the surface. They range in size from the large ones found in the lateral portions of the abdominal tract to the small tufts resembling true down. The ruffed grouse is well known for its large aftershafts. This is a second shaft bearing most of the typical feather parts and attached near the base of the main shaft. Thus the feather is more or less double. In this species it is found on all of the semiplumes and the teleop- tiles, except the rectrices and functional remiges. In the highly modified feathers and some of the coverts, it often cannot be distinguished. In all cases it is entirely plumulaceous or downy and coloring occurs only at the tip. Usually the aftershaft is one-half to three-quarters the length of the main shaft. Numbers. The feathers of a female ruffed grouse* were counted as they were plucked. The results are shown in table 101. The figures, however, represent only the number of teleoptiles and semiplumes present. It is to be noted also that in some ca^es the divisions between the regions of the tracts of necessity had to be arbitrary. TABI.K 101. NUMBER OF KMVniEHS' COUNTED ON AN INDIVIDUAL lUJFFED GROUSE Troct Numbr^r ('aplLuI 776 Spiiiiil I'*7 Posterior :uio Oil CAaiui It Cuudul i(>t Vcntml Crrvinil. Siili-miilnr. Intnr-ramnI 2U 158 lOfi I 11 Alar 2a 1 Crurul uiid Pedal 710 Total 4.3<12 ♦Includes only tcleoplilcs and somiplumns. i.r^. all llip fi-jitlii-rs I)(i|nil;irly thoiitjhl (if ;is surh. Sexual variations in the number of feathers are minor. For instance, in the cervical region the total number is essentially constant, although a greater number are modified to make up the ruff in the male. Similarly, the occasional bird with more than eighteen tail feathers is more frequently a male. In many other birds, Wetmore has found small differences between the sexes in the total number of feathers and probably the same is true of this species but little significance can be attached to such variations. In fact they are probably more closely associated with size than with sex. Seasonal variations in the total number of feathers have not been studied in the ruffed grouse, although Wetmore has found variation of this kind in a number of other species. j)ar- ticularly among the small perching birds. Following the normal fall moult, most of the feathers are immediately replaced but others do not grow until the approach of winter. Simi- larly, during the spring, as the temperature increases, the number is gradually decreased by shedding, a loss which is not replaced until the normal moult or later. Plumage Wear. Plumage wear is not readily evident until several months after the com- pletion of the moult in September, the first sign appearing in the throat region. The gradual fading and soiling of the plumage is not very evident until winter. It is most pronounced in « Taken near Ithaca, N. Y., April 27. 1937, weight 539.7 gm. lii THE I'TKR] I.OGR.II'in Ol' THE lU EEEl) GROUSE llio alidomirKil r(j;i(iii ulirn- llic wliili- limis Id a sooty gray and in tlie lighter regions of the head. But occasioiialK the |)himaf:e is rcmarkabl\ clean at the end of the summer when the next moult takes jjlacc. Kiivironmental conditions vary so widcl), however, that this wear may differ considerably between localities. Age Variations. Age variations are primarily associated with growth. .At hatching, the chick is covered with down. Within the first day, the (light feathers of the juvenile plumage begin to grow and within three or four weeks the body is fairly well protected by the juvenile feathers. With the possible exception of the remiges, which begin to grow almost innnedi- ately after hatching, it is believed that all the juvenile feathers are preceded by down. The distribution of the down is the same as that of the later plumages with the exception of the changes due to body growth. In the down plumage and juvenile plumage the spaces are, for the most part, evident but are smaller in proportion. Concomitant with the growth of the body, the juvenile plumage presents a gradual spreading out of the tracts and spaces. Snowshoes. The "snowshoes", so called, of the ruffed grouse are a uiiicpic temporary grow'th along the sides of the toes, acquired during the fall and shed the following spring. As the name indicates, they increase the surface of the feet I from two to three times I and thereby reduce sinking into the snow. They are included under the heading of feathers since they are derived from the skin and are moulted regularly. They consist of a series of small cuti- cular rods which, in New York State birds, average about 2mni in length*. Growth normally begins about the middle of September and continues for about three weeks. They are usually lost between the latter half of April and the third week of May, although the shedding may begin as early as March. The rods are shed by breaking off at iheir bases in an irregular order, but, more often, first at the base of the toes. * Uttal'**' lias rc[iorIf(i these l»» he whut longer nonhern anil western siihspecies. PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE* By William H. Long Commenting on some of the causes for variation in the behavior and distribution of wild animals living under natural conditions, Charles Elton, the well-known English ecologist. states''^: "most animals are, in practice, limited in their direct distribution by their habits and reactions, the latter being so adjusted that they choose places to live in which are suit- able to their particular physiological requirements . . . every animal has a certain range of external conditions in which it can live successfully, ultimate limits of the environ- ment are set by its physiological make-up, if these limits are reached the animal will die." A knowledge of the physiological limits of tolerance and susceptibility to normal and abnormal influences is an important objective in wildlife research. Yet the physiology and physiological needs of wild animals, particularly upland game birds, are practically unknown. Therefore, a study of the interaction of vital processes and environmental conditions, as well as of the role which specific factors or cf)mi)iiiati(>ns of factors such as temperature, food and water play in regulating the distribution and beha\ ior of a species, is valuable as a supple- ment to ecological investigation. Through direct or indirect effect on the internal \ital processes and external physical reac- tions of an animal, variations in air temperature, air movement, radiation, evaporation, humid- ity and light govern to a large extent the daily and seasonal activity of the animal in its nat- ural habitat. When unfavorable conditions exist, physiological resistance and mortality increase. On the other hand, when favorable living conditions are maintained, both mortality and physiological resistance decrease, with the result that wild game species have a better opportunity to survive, reproduce, and increase in number. An individual bird is not an entity isolated from the habitat. Rather, there is a continual exchange of energy between it and the environment. The habitat, in the form of vegetative. cover and its by-products (food, minerals, water) furnishes the potential energy. The bird, through processes of internal metabolism, absorbs and assimilates the energy, the waste material (carbon dioxide, water, heat, etc.) being returned to the environment. If the habitat does not furnish an adequate supply of potential energy to meet daily requirements for maintenance of physical activity and vital processes, and suflTicient protection against adverse weather conditions and animals of prey, then the inherited ability of the bird to support bodily functions at normal levels is weakened and survival time is shortened. When there exists a condition of stable physiological equilibrium with the habitat, the bird will be ener- getic and better able to resist unfavorable change. When we know more exactly how much of the behavior of a game species is based on the interaction between vital processes and environmental conditions and how each physical factor or combination of factors regulates internal and external activities, including the utilization of food and water, then the game manager will be far better equipped to handle important wildlife problems. * Tlie studiL's iiiMiii Hhicli this paper is basrd wrrp I'onilucU'd by the author in the Physiological Wildlite Research Labora- tory located at Cornell University. Ithaca. New York, and established jointly by the New York State Conservation Department, the United States Fish .ind Wildlife Service, and the New York State College of Agriculture. Acknowledgment is due Cuilford\ surface or by ])lacing the bird in a cage and obtain- ing a graphic rec-ording of the res|)irator\ niovcTncnt 1)\ means of the k\ mograpli. Records of iuail beat were secured li\ using a sensitive stethoscope and s\ nc hronizing the sound with "hand-tapping" a ])encil point cm mci\ing pa|)cM- or the |irc-ssing of an auto- matic electric counter. Various special ici hnicpics were devised ij\ ihc aiiliioi to iiuct specific needs as the work proceeded. In |)lanning the studies to be condncted emphasis was placed on securing information per- taining to the general plnsiological characteristics of the species which might ha\e a bearing on management practices as well as to securing basic data* prerequisite to conducting more ♦ UnforciinaCely urmrc ilurn not [icnnil llie incluitiiin of niuiiy artuils. liolli i\tlli rrtp.-.t lo .lata ami In liniiiucit rni|ili)>r.l, wliicli have been invoUeil in thii project. PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 751 elaborate physiologic investigation on grouse. Therefore, attention was directed chiefly to those physiological activities which are intimately associated with the production and main- tenance of body heat, namely, body temperature, breathing and heart rate; ingestion and digestion of food and water; and change in body weight. Experimental birds were supplied from hand-raised stock at the Research Center. At the laboratory, when not on experiment, they were held in a tem|)erature-regulated room under natural lighting and were fed the usual ration employed at the Center. One must bear in mind, however, that there are undoubtedly some differences between the reactions of such birds and those of wild grouse. Determination of Body Temperature and Respiration Rate Experiment 1 dealt with the determination of the average body temperature and respiratTon rate of adult grouse under conditions of normal cage acli\ itv. Cunnilative data from ten birds were secured on consecutive days for several weeks under identical environmental con- ditions. The results are presented in table 102. TABI-E 102. AVEIUCK MKCTM. TI;MI>1:H \Ti;nK AND HRSPIHATION IVVTK OF. ADULT GHOUSi:* Sex Air temiicrature Time of (lay NiimlMT of obsorvalinns Rectal temperatuFG Respiration per minute birds Avorago Standard error Average Stjiiidard error 5 5 5 5 5 5 male IViiialr riijilc l.-„ial.' niixivl mixfid M'F. ().') (i.'i HO-H.i 80-85 inorriiti^' iiinniiiif; urtxTiKKiii iirt4'riu>uii morning' nflcriKH)!! 12.-! lO.'l 7') 17 60 41 107.26">K. Iti().>)'( i()7.().'; 1(17.7(1 • 108.24 108.32 0.03.'; o.n.i'i 0.040 0.087 0.198 0.155 64.5 58.2 67.1 59.8 71.6 74.0 0.871 0.969 i.lin 1 .'too .1.140 4.640 ^Normal cage activity. Comparison of the average rectal temperature and res|>iiatiiiM lalc iridiiates some signifi- cant biological difTcrences lietween sexes. The morning |pni|)crature and respiration rate of the male grouse were higiicr than those of the females, in tiie afternoon res])iration rate was higher for the males but both sexes were (juilc similar as to temperature. There was a consistent rise among indi\iduals of the same sex between average morning and afternoon temperatures, indicating that tlie higher afternoon level represents a real physiological difference. A similar relationship was also noted between rectal temperature averages secured at 6.5°F. and 85°F. In this case statistical evaluation of the data indicates a direct effect of air tem])erature itself upon metabolic processes. Diurnal Trend in Body Temperature Experiment 2 was undertaken to establish the diurnal trend in body temperature. Two groups of birds were used, each com|jrising five male and five female grouse. Rectal tem- peratures were taken for each group at regular 3-hour intervals for two separate 24-hour periods beginning in one case at 8:30 a. m.. in the other at 10:00 a.m. This procedure per- mitted plotting the data at intervals of 1 Vl' hours. With one exception (the factor of light) both groups were subjected to comparable conditions of environmental temperature, activity and food. With respect to light. Group A was exposed to an additional hour of illumination at night. 752 PHYSIOLOGICAL STVOIFS OF THE RVFFFJ) GROiSE The results, summarized in figures 83 and 84, clearly show the existence ol a diurnal variation in rectal temperature. iriHicalinji a similar trend in body temperature. Highest "F. 108 5, "F. ■ 106.5 1080 107.5 UJ a- D (- < a: 1070 Ld Q. UJ I- _l 1065 < I- o LJ o: 1060 I05.5L 105.0 — Group A — Group B _!_ 1060 107.5 1070 10 6.5 106.0 105.5 U 105.0 8 30 11:30 - -A.M. »K- 230 5 30 8 30 P. M . ll;30 4^ 230 5:30 - A.M.- 6 30 TIME OF DAY I III KK 83. DIURNAL TRENDS IN RECTAL TEMPERATURE OF TWO CROUPS OF ADULT GROUSE, CROUP A OF WHUH \V\S EXPOSED TO W \nr)nin\\l, IIOIR OF II.I.IMIN VTION \T MCHT temperatures were reached between 2:30 and 5:30 i).iii.. and lowest lictwcen 11:30 |).tn. and 2:30 a.m. The average difTerotup helween the hiirhesi and lowest leiiiperaliires dining the 21-hour period, was: For males: 2.69°F.± 0.257" (standard error ) For females: 2.29°F. ± 0.139° (standard error 1 With reference to the efT<'cl of light on the diurnal curve, the peak in l)<>d\ tem|)erature reached at 5:30 ]).m. for (".roup A seems to have been due chiefly to extra activity and feed- ing; allowed them during the iiddilional hoiii of light. Effect of F.xcitement and Activity on Ut)u\ Temperati hk Experiment 3 was designed lo measuie the eifi 1 I nf ernolinMal <\eili'nieiil aiMl Mm>eiil;n ac- tivitv on rectal lernpi'rature and rale of breathing. Four fcriude and l«o male birds, in a state of active digestion, were used. They were renioxed frmn llie lage one al a lime fio body temperature and respiration nx-ordings. Kach wa> llien put into aiiolher (age ami PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 753 106.5 lO&O 105.5 J '-'105.0 TIME OF DAY FICURK z 2 65 a: 111 a. fiO Ld 1- < cc 55 50 RECTAL TEMPERATURE Be+ore txciienrten+ RESPIRATION After E.>:ci+cmen4 FUUIRE o.S. EFFECT OF EXCITEMENT ON RECTAL TEMPERATURE AND RESPIRATION RATE OF ADULT CROI SE 754 P/nf^lOLOGIC IL STl DIES OF THE Rl FEED GROUSE fdrccri t(p run and mkim' ;ili(nil fni ten niituilt'>. aftiT which temperature and respiration were again recorded. Ihe results of this test are sliown in figure S5. Thev indicate a significant increase in both respiration rate and body temperature foUowing ten minutes of activity. Ekfkct of Rainfai.i, on Maintenance of Body Temperatire Experiment I itnolved a stud\ of the effect of rainfall under various conditions of air temperature and air movement on the bird's al)ilil\ tu maintain its body temperature. Two groups (A and C) of five grouse each, and one 1 15 I of six. were used. All were in a condi- tion of active digestion during the tests. Rainfall was simulated in the experimental cabinet by spraying water on the birds. Air movement was slow. Specific conditions for each group were as follows: Group A — Exposed to Air Temperature 40° F.; Humidity, GO'^'i ; Water Temperature 44°F. Group B — Exposed to Air Temperature 65°F.; Humidity, SO'i; Water Temperature 44°F. Group C — Exposed to Air Temperature 80°F.; Humidity, 35%; Water Temperature 62°F. The results are shown in figure of). It is to be noted that in each case a sharp drop in body temperature occurred during the half hour inunediatel) after the birds were wet. This 0 20 40 60 80 100 120 140 160 180 200 220 2*0 280 280 300 DURATION OF TEST IN MINUTES FIGIRE 86. TRENDS IN RECTAI, TEMPERATIRE FOLLOW IM. \\ K TTING AMONC, THREE (;R01PS OF ADULT GROUSE EXPOSED TO DIFFERENT CONDITIONS OF AIR TEMPERATURE AND HUMIDITY was most severe and recoverv was slowest among Group A. which were exposed to a water temperature of 14 F. at an air temperature of 4fl°F. Even at an air temperature of 80°F. welting caused a marked and rapid fall in \un]\ temperature (Group C). The data clearly show that wetting the binK willi u.ilir is a factor of considerable ])hysiological importance in itself. The cdndiinatinn mI I.iu air temperature anlill more ef- fective sinie the rate nf (liii|i in IhkK tempcralnre w'as faster and rate of recovery slower. I'nder conditions such as those to which Groups A and H were subjected, continued exposure without good bodiK protection would cause death. PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 755 In another part of the experiment five apparently healthy grouse were subjected to similar rainfall at an air temperature of 62°F. and an air movement of 15 miles per hour. Figure 87 shows that only two of the test birds (Nos. 8 and 29) withstood this treatment and sur- •F. loe 106 tr D H < Q. 5 u H _j 100 < I- o tu o: 9a 96 94 40 60 80 100 120 140 160 DURATION OF TEST IN MINUTES riGlIRK 87. TRENDS IN RECTAL TEMPER\TIRE FOLLOW I.\(. WETTLNU AMONG DISEASED AS COM- PARED WITH HEALTHY ADULT GROUSE vived, whereas grouse No. 13 died within four hours after the completion of the test, its body temperature falling to a very low level. Grouse No. 37 died two days after the test and grouse No. 21, three days after its completion. Postmortem examinations indicated that all three of these birds were suffering from enteric disease*, although prior to the test they appeared healthy. These data indicate that pathological conditions such as enteritis or in- testinal gout when latent or inactive in the body of a grouse can he activated by a drop in body temperature as a result of wetting the body with water of chilling temperature. Heart Activity as an Index of Vitality Experiment 5 consisted of a series of tests to determine the significance of heart activity as an index of vitality. To begin with, a general study was undertaken of the relationship between body temperature, respiration and heart rate in relation to activity, inactivity and emotional excitement. Grouse in different groups were held inactive from IV2 to 2-^$ hours. * Diagnosed by ConBervdiion Palhologisl, Dr. Frans C. Coble at the \('ildlife Research Center. Delmar. New York. 756 PHYSIOLOGICAL STUDIES OF THE RIFFED GROUSE Previous to testing some of the birds had been living at 65°F. and 80°F. Some were meas- ured immediately after ingesting food, others a few hours after eating and still others after fasting from 100 to 148 hours. After the birds had been held inactive for the time specified, they were subjected to a short period of activity, following which measurements were again recorded to determine the influ- ence of this type of activity and excitement upon physiologic factors. The resultant data are presented in table 103. TABLK 103. EFFECT OF IiWCTIVlTV ON THE BODY TKMI'EH XTIHK. HEVKT R\TK AND RESPIRATION OF BOTH FEEDING AND FASTINti GROUSE Condition of birds Number of birds Air temperature Time of inactivity Measurements recorded At start After inactivity Drop After excitement Rise Average soon after eating Fasting 100 to 148 hours. Normally feeding Average soon after eating Fasting 100 to 1(8 hours. Normally feeding Average soon after eating Fasting 100 to 148 hours. 6 6 {'I 6 6 {'I 53°F. 63 63 80 63 65 65 63 63 2H hrs. l-S/6 2% 2\4 1-5/6 2H 2i4 1-5/6 108.2°F. 106.9 107.7 108.4 73 53 64 75 Rectal temperature 106.1°F 10S.8 106.6 106.2 2.1">F. 1.1 1.1 107.0°F. 106.8 107.3 107.3 Respiration rat^' — p^'r minute 51 46 46 54 18 21 79 61 67 78 Heart rate — iicr mioute 333 310 280 237 53 73 349 334 0.9° F. 1.0 0.7 1.1 28 IS 21 21 69 97 Both inactivit} and excitement liad a signilicant effect upon heart rale and body tempera- ture. This rapid physiologic adjustment to changing environmental conditions demonstrates considerable lability in heart beat in whirh the great increase from the minimum level of repose to the level of emotional excitement is particularly noteworthy, especially for fasting grouse. Because of indi\iilual variation in the frequencN. (Iiiratioii and intcnsitv of active periods, it was not practical to plot the average trends for the various factors for all grouse used in this experiment. To do this would definitely mask the true effect of repose on metabolic ac- tivity. However, the physiologic reaction of two grouse to repose has been plotted at four- niinule observational intervals (figure 88). These records are representative of ten ntlu-r individuals. There was a considerable reduction in rale of breathing and heart beat and in body tem- perature as these grouse became more and more inactive. Of the two iiirds used, liie female exhibited a greater rate of reduction in the intensity of the three physiological activities. The metabolic intensity of the male remained higher and required a longer time to reach a basal level at about the 2 1 6th mimile than the female whose basal level was reached at about the 168th minute of slud\. .Slight movements of the external body (iiidiralcd li\ \ in figure 88) resulted in a small increase in the several physiologic activities. Of interest are the observations, designated by ® at various points in the figure, that a slight rise in body temperature preceded a drop of greater degree. This phenomenon was noted fre- PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 757 quently throughout the tests. It apparently is not a result of visible muscular activity for the birds were quiet when the change occurred. More probably it is caused by cellular varia- tions in heat production and the subsequent flow and distribution of heat throughout all parts of the body. RR 72 ^ KR 324 H R - Heart Rate R R - R«apiratior^ Rate A - Activity Cnoturat) EXCITED-Artificolty Stimulated 2 107 0 I go 100 120 140 160 leo DURATION OF TEST IN MINUTES 220 0 20 40? 60 ii(;iKE f>R. piivsioLOGie reactions ok two <;k()1 se to kepose The rate of metabolic activity was more rapid at the beginning of the test, but as a basal condition was approached the rate decreased. When final basal levels were reached both of the grouse (representative of ten individuals! were removed from the "holding board" (figure 89) and excited emotionally and muscularly. The result of this stinmlation was quickly reflected by pronounced increases in body temperature, respiration and heart rate. These results again clearly demonstrate the intimate relationship between environmental con- ditions and physiological phenomena. Next the rate of heart beat was established fm Imlli feeding and fasting grouse at different air temperatures. In these tests the heart rate was classified under three headings, that is, (1) the standard rate of the inactive bird held for studv immediately after remo\al from its cage; (2) the standard quiet rate of the inactive liird held quietly for some time after re- cording rate No. 1; and (3) the excited rate resulting from muscular stinndation. The air temperature indicated is that to which the birds were accustomed to living before testing. The results for feeding birds are presented in table 104. Throughout these tests the heart rate of birds with food was higher at 40°F. than at 65°F. Immediately upon removal from the cage, female grouse exhibited a faster heart rate than males at both OST. and 40^"F. As indicated by the standard deviation the range of varia- bility among both sexes was considerable. In all observations the lowest range for females was approximately equivalent to the highest range for males. Likewise, after the birds had had an opportunity to quiet down, the same relationships existed. 758 PHYSIOLOGICAL STUDIES OF THE RIFFED GROUSE 2 z X u o o o X PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 759 Data recorded when the birds were excited showed no appreciable difference between the sexes. The extreme variability of the heart rate under these conditions, however, might easily mask the difference indicated in inactive and quiet birds. TABLE lot. HEART RATES OF ADULT GROUSE LIVING* .\T TWO AIR TEMPERATURES WITH RESPECT TO DIFFERENT DEGREES OF ACTIVITY .Sp, Air temperature Measurements reoordedt Number of birdsA Average Standard deviation Standard error Standard inactive rat* 24 18 .-(6 IH male male female female 65° K. 40 65 40 306.S 321.6 366.7 379.4 36.3 36.2 25.9 30.6 7.6 12.5 4.3 10.6 Standard quiet inactive rati* 24 IB :i6 18 iiifili' male female female 65 40 65 40 295.5 308.5 3S8.S 367.6 32.5 34.7 23.6 2T.9 12..1 15.5 liiJ Standard eiriled rate 24 18 36 18 mule male female female 65 40 65 40 386.6 .191.3 399.3 416.2 .36.1 33.8 12.7 33.7 13.6 15.1 3.9 15.0 *Normal cage life. AComposed of 288 individual records, tKeats per minute. When the data for quiet and excited birds are compared a tremendous and almost instan- taneous increase in heart beat under stress of excitement is evident. To determine the influence of food on the rate of the various physiological activities the birds were subjected to fasting for 100 hours and then tested (table 105). TABLE 105. AVERAGE STANDARD QUIET HEART R.VIE OF INACTIVE ADULT GROUSE 100 HOURS AFTER WITHDR.\WAL OF FOOD Number birds Sex Air temperature Average heart rate Standard deviation Standard error 9 9 9 9 Male Male Female Female 6S'F. 40 65 40 295.6 282.0 308.0 294.0 24.9 25.3 43.6 20.0 8.9 8.9 15.5 7.1 There was no significant difference between male and female heart rates after fasting 100 hours. At this stage of fasting the destruction of cellular tissue and fat deposits was progressing rapidly, the bird probably having utilized most i>f it.* reserve supph of liver and muscle glycogen. Practically all of a grouse's energy for maintenance of vital processes is derived from the combustion of body fat after fasting only 36 to 48 hours. Thus the true sex difference in heart rate is masked by the catabolic activities being carried on at an ac- celerated rate in the body. It should also be noted that, in addition to being considerably slower and showing less variability, the heart rate of fasting birds was lower at 40°F. than at 65°F. It is apparent that cardiac action is affected by fasting in similar physiologic ways 7f)0 PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE as body tfiiijx'raliiic and rale uf l)reathinj;. llic prcal iiifgularilN of heart aclinn and llii' vital capacit) of the heart to coin])eiisatP for adverse environmental change, is indicated by the standard deviation with respect to the \ariabilitv of the rate of heart beat. In these tests of grouse the number of heart beats per minute I sexes combined I averaged 342.6 and 350.5 at 65°F. and 40°F. respectively for birds on food, and 301.8 and 288.0 re- spectivelv for fasting individuals. Statistical methods described by Baten^ for calculating data were used to measure the vital relationship between the .physiologic variables where x = heart rate per minute; y = body temperature; and z = respiration rate per minute. By this means the significance of the rela- tionship and the amount of dependence each variable has upon one or more of the others can be demonstrated. Calculations of the various coefficients"'*" for both feeding and fasting grouse are presented in tables 106 and 107. T\RLK 106. CALCULATION OF AMOUNT OF DEPENDENCE BETWEEN PHYSIOLOGICAL REACTIONS OF FEEDING ADULT GROUSE* Reaction Number of observations Average rate Standard error Correlation e(ricient Heart rate (x) 27 27 27 301.2 107.1° K. 301.2 .19 7 S.2 0.2 5.2 1.9 0.2 1.9 Rxy = +0.58 + 0.133 Body t«mperature(y) Rxz - +0.48 + 0.108 27 107.1° K. 27 59.7 Ryz 1= +0.6B ±0.154 ♦Average cage activity. TABLE 107 CALCULATION OF AMOUNT OF DEPENDENCE BETWEEN PHYSIOLO(ilC\L REACTIONS OF FASTING ADULT GROUSE* llf'fiction Number of observations Average rate Standard Correlation error coetlicieut 18 18 III 264.4 106.3° F. 2h4 4 11.8 lUy = +0.60 + 0.150 0.3 118 Hiz = 4-0 72 + n 120 111 1 IB.O 2.7 111 ; 106.:t° K. Ill 48.0 0.3 2.7 Ryz = +0.41 ±0.208 *Average cage activity. ("omparisoii of the differences between averages for the three physiologic reactions for both fasting and feeding birds indicates the following t-values of significance: 2.63 — Heart rate 2.07 — Body temperature 3.59- Respiration rate These data support our biological understanding of the intimate iiiterplav of these physio- logic factors. Since the coefficients are all positive and highly significant, either when related individually to each other or when all are related together it niav be stated that a higher heart rate is intimately related to a higher body temperature. Thus when body temperature increases PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 761 or decreases, both rale of heart and breathing are accelerated or retarded. Because of the great degree of interdependence among these physiologic variables* it may be safely concluded that rate of heart beat per minute and body temperature are fundamentally significant as in- dices of the level of vitality. Respiration rate per minute is a factor of less importance as an index of metabolism, especially for the type of experiment described here. Under other experimental circumstances, for example where an excessive change in body temperature oc- curred, there might be pronounced changes in rate of breathing. Resistance to Starvation Experiment 6 was devised for measuring the resistance of ruffed grouse to starvation using rectal temperature and respiration rate as indices. First, all food except water was withdrawn over a 10-day period from 12 grouse, six of which (Group A) were held at 6.5°F. and six (Group B) at 40°F. throughout the test. Records of rectal temperature and respiration were taken three times daily — morning, afternoon and midnight. A summation of the data for each of these time intervals is presented in table 10}'. TABLE 108. EFFECT OF STARVATION WITH WATER AVAILABLE UPON Till'; KICCTVl TEMPERATURE OF ADULT GROUSE Number of birds Sex Time of day llcctol tcmperaluiv Group Average Standard Stuudurd deviation error (iroup A, at air tcmperutiirc 65° F. 3 3 mole female morniug afternoon midnight morning afternoon midnight 106,37"'F, 106,25 103,51 106,56 106,70 103,56 0.352 0.410 0.303 0,128 0.434 0,789 0.117 0.1.37 0.114 0.0 13 0.14S 0.299 Group B, at air tcmporattire 40" F. 3 3 mule female morning afternoon midnight morniug afterncxin midnigbt 105,53 106,17 103,96 inh,oi lll.'..Kh 103.90 0,482 0,265 0,830 1.114 1.480 0.316 0,161 0.088 0.339 0.390 0.406 0.129 Of interest to the game manager is the greater drop in the level of rectal temperature for fasting birds at midnight compared with that of well-fed birds at the same hour ( Experiment 2, figures 83 and 84). During the night there is a natural rhythmical lowering of metabolic activity and external physical movement. Also between midnight and dawn under outdoor conditions the lowest air temperatures usually occur. As the air becomes colder, however, the metabolic rate would tend to increase in order to prevent a drop in body temperature. Under such conditions a fasting bird would have greater difficulty in preventing an excessive reduction of body temperature than would a well-fed bird. The difference between the averages of data obtained at 40°F. and 65°F. shows that there was a high physiological correlation between body temperature and environmental tempera- ture at these levels. The smaller standard error for females at 65°F. indicates less variability in their temperature than in that of males at the same air temperature. At 40° F. the degree of variability apparently was not significant between the sexes. * As indicated by the highly BigniBcant, positive multiple cone + 0.60 ± 0.165 762 PHYSIOLOGICAL STIDIFS OF THE KLFFED GROISE The average trend of rectal temperature during the period of the test is plotted in figure 90. It shows greater variability among the birds living at 65°F. (Group A). Also the birds in this group maintained their temperature at a higher level than did those in Group B which were subject to greater loss of body heat by radiation at 40°F. than at 65°F. It is probable that a difference of 20 degrees in air temperature would have a more pronounced effect upon the metabolism of fasting grouse than upon that of well-fed grouse. 107.0 I 106.0 1 a. I- < q: I0S.0 104.0 < U UJ Q^ 103.0 102.0 '-' L, Of-oup A at <<>b"r Group B ol 40 T _l_ -L. 5 2 2 2 < a. 5 < 72 96 5 Z 0. 5 z 2 2 2 < CL 144 ' r I 10 7 0 FI(;l RE 'JO. 120 144 168 192 216 TIME OF DAY & HOURS OF FASTING TKKND.S 1\ KKCTAI, TK M PKRATl'Kl-: .\MON(, TWO GKOl PS OK K.V.STING ADl l.T (.KOI SIC MKl.l) \l l)IFFER^:^T AIR TEMPERATURES The respiration rates recorded for these birds are sununari/.ed in table 109. When com- pared with ihe data secured for well-fed birds in Experiment 1. tabli' 102. they show^ that fasting lowered this rate in both sexes. Also shown is a distinct difference in this reaction at 40°F. and at f)S°P'. similar to the trend in rectal temperature. Furthermore, tlie difference between the sexes at 40° F.. namely 13 respirations per minute higher for female, is statis- tically significant. TAHLli 109. liKKECT OF STARVATION UPON HESPIK.VITDN WVVK OK ADIJI.T <;H()I SK* Si-x Air lriii|H'ratiiri> 1 J UfMpirutiuii rale P.-riiKl , Standard Average deviation SUndaril error A.M. to A.M. A.M. to A.M. A.M. U> A.M. A.M. to A.M. Female Female Male Mule 65° F. 40 65 40 55.0S 3.08 47.34 3.79 56.34 3.99 60.49 4.36 1.07 2.25 1.40 1.49 *Waler available. As a second part of this experiment, six additional grouse I Group Cl were held at 65°F. with neither food nor drinking water available for 84 hours. The average rectal tempera- tures and respiration rales of these birds are presented in table 110. During the period of study half the birds died. Since this group wa.* tested under identical environmental condi- PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 763 tions with the other two, except for the absence of drinking water, it would appear that such water, or a suitable substitute, is a daily physiological need of grouse. TABLE 110. EFFECT OF STARVATION WITHOUT WATER UPON THE RECTAL TEMPERATURE AND RESPIRATION R.VTE OF ADULT GROUSE* Measurements rerded Numbpr oi Sex Time (if (lay hirdu Average SUndard Standard deviation error H Petal temperature :i Mill.- Morning' Aft^rruxm l()7.1<)° F. n.B29 ().8T0 0.479 0.435 :i FemalR MoniiiiK Afternoon IIIO.TI Klh.tH 0.8<)6 O.ihV 0.H8 0.235 3 Male MorniuK Afternoun 62.5 .56..') Heapiration rate 6.02 4.15 3.47 2.08 .1 Keinult* VforiiiiiK Aftern(Kni 57.0 59..1 5..'j:l 3.16 2.66 1.58 *Group C, at air temperature 65° F. The influence of length of fasting, time of day, and absence of drinking water upon the leinpt-rature-regulating niechaiiisin is presented in figure 91. The female grouse in Group C showed least resistance to fasting, as indicated by the trend of their temperature, especially at midday. After the 48th hour of fasting without water neither the males nor females were ioe.O| 107.5 Id D 107.0 !< u Q. IS 10 &S UJ I- _l < I- 10 6.0 1 U 10 5.5 ,106.0 107.5 107.0 Ji0&5 106.0 105.5 105.0 0 24 4a 72 9 6 NOON P.M. NOON P.M. NOON PM. NOON P.M. NOON U 105.0 DURATION OF TEST IN HOURS FICURK 91. TRENDS IN RECTAL TEMPERATURE AMONG MALE AND FEMALE ADULT CROU.SE FASTING WITHOUT DRINKING WATER 764 I'lnSIOLOCICAL STL DIES OF THE RL FEED CROiSE able to iiKiease their mctaliolisin snllicii-ntly to prpveiit a hrpakdowii of their temperature- regulating mechanism. In this i)arti(ular instance there was a significant rlilTerence hetween the averages of body temperature for males and females, beginning at the 'Mtih hour of fasting. In table 111 the body temperatures of Groups A and C are compared with those of nor- mally feeding birds. The rectal temperature of Group C birds at the 84th hour of fasting averaged higher for both sexes than for Group A but lower than for individuals that were not subjected to fasting during these tests. The experimental data from this group demon- strate the effect of both active digestion and starvation upon body temperature. Variability in rectal temperature, as indicated by the magnitude of the standard error, was least for the control birds and most for those individuals fasting without water. TABl.K 111. COMPARISON OF BODY TEMPERATURE OF FEEDIN(; GROUSE WITH THAT OF GROUSE AFTER FASTINC; lil Mot US BOTH WITH AND WITHOUT WATER Number of birds Sex Rectal temperature (Jroiip lemiwrature of day Average Standard error 10 6 6 mixed mixed mixed 65° F. 65 65 inorniiiK afUTiioou luoniinK 107.14° F. 107. 6<» 10(1.47 0.0.'i4 0.044 0.080 afternoon 106.48 0.141 0.46.1 afternoon 106.73 0.33S ♦Average caj,'*' activity. ^Group A. t(iroup C. When fasting without water, grouse apparently are subjected to rapid dehydration of body tissues. The drying-up process is of such magnitude as to cause a definite increase in mor- tality. The data from all groups of birds in these tests clearly indicate that grouse have definite daily need for water to satisfy particular physiologic functions. Due to limited data the actual cause of carlv death of grouse fasting without water is not clearly understood at this time. Effect of Fasting on Body Weight Experiment 7 was supplementary to the preceding sluch and irn(il\td cLila taken coinci- dental!) from the same groups of birds. Its |)urpose was to stndx liie elTcil nf fasting I both with and without water) upon body weight. By subjecting the birds to fasting conditions the resultant data fnrnisiicd a fundamental base-line of minimum vital needs for maintenance of the various physiologic functions. By superimposing upon the base-line nf minimum needs the effect of such factors as muscular activity, digestion of food, and air temperature, the maximum phvsiologic needs for main- taining \ital functions at normal levels will be revealed. The 18 birds were subjected to prolonged fasting as lollcius: Group A — 6 grouse at 65°F.. withdut food biil with water. Group B — f) grouse at lO'F.. without food but with water Group C — 6 grouse at 6.S°F., without food or water PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 765 111 each group, tin- scxr.s witc oveiily ilividfd. The results (if the total |)erif)d of fast are recorded in table 1 1 2. The ability to withstand fasting does not seem to be entirely dependent upon nor strictly proportional to body size (weight) alone. Temperature factors appear to have been highly significant in the case of the three female grouse which lost approximately .31 per cent of their weight after fasting 189 hours at 40°F. The trend of daily weight change for individual fasting grouse in all groups is shown in figure 92. Figure 93 shows the percentage of weight loss. The slopes of these curves are interesting in several respects. They indicate that (1) the daily ingestion of water pro- longed life and sustained body weight; (2) grouse of lower weight suffered a more rapid loss in weight*; (3) heavier grouse had a tendency to maintain a more constant weight loss: and (4) percentage losses in weight were more or less proportional to the hour of fasting. Sixty-seven per cent of the grouse in Group C, on test without food or water at 65°F., died, on the average, at the 88th hour of fast, with an average loss of 16..5 per cent of the initial weight. When this weight loss for Group C is compared with the average percentage loss for Groups A and B at the same hour of fast, it was revealed that the latter had lost only 11.7 per cent of their initial weight. This small difference is highly significant since the 88th hour of fasting with water represents only 40 jier cent of the total period of fasting to which they were exposed. Undoubtedly this greater degree of resistance to fasting was due to the ingestion of water by those birds that were allowed it during the test. The average distribution in weight loss ])er 24 hours showed the rate to be considerablv less during the daytime than during the night, the greatest loss occurring from midnight to 8:00 to 9:00 a.m. TABLE 112. LOSS IN WEIGHT AMONG ADULT GROUSK DURING STARV.VTION ILNDICH DIFFERENT CONDITIONS Group Number of birds Sex Air temperature Percent humidity Initial weight Final weight Percent weight loss Length of fast Loss per hour Mortality Kosting, Willi water (Group A) 3 3 male female 65° F. 65 30 30 613.3 gm. 533.3 468.7 gm. 411.0 23.6 22.9 234.7 hrs. 232.0 0.62 gm. 0.53 2 Fasting, with water (Group H) 3 3 male female K) 65 (>5 602. TRENDS OF 24-HOUR CHANCES IN BODY WEICHT OF ADULT GROUSE DURING PROI.ONCEn F\STIN'C. WITH AM) WITHOUT WATER Analysis of data from 16 observations on the relationship between water consumption anil change in body weight were calculated statistically as follows:'" Predicting equation: V 103.25 \X)H^^\ (Correlation coenicicnl: Ixxy -= —0.65 ± 0.15-1 I standard error! Since the \aliic of llic ^/atd- o+ C»5** ■ Group B , fa2.ting wi+h wat«r a+ -40* • Group C jfoa»+ir)Q without vyaterat Ci5* 24 48 72 96 120 144 168 192 DURATION OF TEST IN HOURS 216 240 ilOO 95 90 S5 80 75 70 65 FIGURK 9,'^. TRENDS OF PF.RCENTA(,K LOSS IN HODV « KI(;HT OK \l)l l.T GROUSE DURING PROLONGED FASTI.\(. WITH \M) WITHOIT WATER the dye in voided excreta. A summation of the data secured is presented in tables 113 and 114. It is apparent that rate of digestion is influenced hy both air temperature and composition of food. At the lower air temperatures digestive activity was appreciably slower. In this connection it is to be noted that the rate of activity of the various digestive enzymes is retarded at lower air temperature, but just how this phenomenon is related to a slower rate of digestion is not clear for the body temperature was not materially affected at the experi- mental air temperature, as shown in tables 113, 114 and 115. When the data are interpreted statistically (table 11,5), it is of considerable interest to note TABLK 11:5. n.vn-: ok digestion of vdult chouse feeding uninterruptedly AT DIFFERENT MR TEMPERATURES Number Sex Air temperature Per cent humidity Ration* Rate of digestion of birds Average Variation 4 4 4 8 4 6 4 mixed female male mixed male mixed mixed 86° K. 80 80 65 65 52 40 57 44 44 30 24 63 60 scratch grain and mash mash scratch grain scratch grain and mash scratch grain scratch grain and mash scratch grain and mash 67 min. 107 140 139 145 239 242 55- 79 min. 105-110 130-150 130-160 135-155 190-288 172-310 ♦All diets irirliidfd ulfalfa, npple and wiiter. 768 PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE the finding of a negative coefficient for birds actively digesting food and a positive coefficient for fasting birds. The values of both coefi'icients are biologically very high and demonstrate a strong relationship between air temperature and digestive activity. I'Mui: 111. n\ri': ok digestion ok adult (jkouse kvteh fasting a.t DIFFERENT AIR TEMPERATURES Number Sex Air temperature Per cent humidity I..ength of fast Rate of digestion of birds Average Variation 1 mixed 82° F. 48 12-1.'. hrs. 75 min. 60- 'HI mill. 4 mixed h.i .10 12-1.-. 90 hd-ll.-. 4 mixed h.'-) 30 78 U6 112-12(1 4 mixed h.'-> .10 192 18.1 16.-.-2I10 4 mixed 40 60 78 102 8.1-1211 4 mixed in 60 192 111 1()0-12.'. T\L5Ll-: 11.5. UEL.VnoNSHll' BETWEEN EN\ IHO.NMENTAL TE.MPER.VTIIVK AND DIGESTIVE ACTIVITY Number of observations Plane of nutrition .\ir temperature Correlation coefficient* 2S 16 feeding fasting 78-100 hrs. 50-86° F. 40-65 Rxy = -0.83 + 0.066 Rxy = -t-0.63 + 0.161 temperature; y = rate of disestion. Physiologically the high negative correlation coefficient is interpreted to mean thai when environmental temperature decreases the length of time required for food to pass through the alimentary canal increases. Under these conditions metabolism would be stimulated by cold but food substances would be retained longer in digestive ])rocesses in order to exhaust and absorb all of the possible available niitriinerits. At high air temperatures the reverse would be the case for actively feeding birds. The high positive coeflicient is of considerable plnsiologic interest. It suggests that expos- ing a fasting grouse to lowering environmental temperature decreases the length of time required for food to digest and be voided as excreta. Greater oxidation of cellular tissue (at from 78 to 100 hours of fasting, essentially a fat metabolism I to combat cold ajiparently stimulates metabolism considerably. .\t higher air temperature the rate of digestion of fast- ing birds would also increase. These two observations on digestive aclivity in relation to air temperature are of con- siderable importance in evaluating the probabilit\ of sur\ival of game species during winter months, especiallv during hours of darkness. Regardless of the temperature effect, the rate of passage of digestible substances through the aliinentar\ c aiial nf the fasting bird was cnnsid- erably faster than for the feeding bird. Rki.ationsiiip of Ckktmn Fmiods to Maintenanck ok I><>l)^ \\'ki(;ht and Vit.ality Kxpcrinient ') iiuoKed llii' lelatimiship nf certain food rations tn tiie inainlenanci' of bod\ weight and \italil\ aimmg adult grouse. I'oods \ar\ ^^ilh respect to their nutritive \alue. If a change occurs in the composition of the food ration. f liPiil miiiiii-ii tti i;iU<" ItlOO grams of water 1°C. 770 PHYSIOLOGICAL STVDIF.S OF THE RIFFKD GROUSE not sufficient to maintain l)odv weight, although \vh\ thev did not eat more is not clear. It is to he noted from previous experiments that increased muscular activity greatly raises the metabolic rate. In confinement, however, muscular activit> is markedly reduced as com- pared with birds whose movements are not greath restricted. Within a certain range of air temperature the metabolism of gallinaceous birds is unaffected by air temiierature itself, tmt either below or above this range it is accelerated to maximum levels. Of the grouse living at 85°F. two (Croup C) gained weight, while five fGroup D) lost. Because less body heat is required as the environmental temperature rises, physiological demands should be satisfied by lower food consumption (caloric values being equal) under such conditions. Yet it is apparent that the caloric intake of the birds in Grouj) D was inade- quate for vital needs. On the other hand. Group C birds consumed relativeK larger amounts of food than was necessary physiologically at an air temperature of o5°F. These two birds were able to gain even more. |)roporlionally. than Group A, although consiiniing no more than (iroup B which could not maintain weight at 65°F. I.SOi Weight Loss Water Consumption m ■ 1.50 125 1.00 Q75 050 025 FEMALE MALE LIVING AT 40° F. FEMALE MALE LIVING AT 65 °F FIGURK '>\. HKI.^TIONSIIIl' HKTVVKKN BODY WKK.IIT LOSS \\l) WATER CONSUMPTION OK M)l l.T GROUSE DURING PROLONGKn KASTIN(. \T TWO MR TKMPF.RATIRKS A possible explanation for \ariations in weight and food consumption is that a tempera- ture of 8.S°F. is physiologically unfavorable during winter months for confined grouse in full |)Iumage and while in a condition oi active digestion. Under these conditions two of the birds manifested polypnea (an exaggerated rate of breathing or panting) which is a chief physical means for cooling the boiK 1>\ tlic vaporization of water from \hr lungs and air sacs. Another phase of the experiment was desigru'd to study the effect of the ingestion of food upon grouse in a physiological state of undernutrition. Ten birds which had survived the PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 771 CO cd H -«! CC H £ o i/3 o a: o Q «! b O O o Q 3s 1 '3 cS O O M ^1 «■] --" « in irtiftM-* s qqeqsq CO o" ro -* II •<3 >> ES t-T3 ti o a c s 2 ©; ^O rO Ov 1 ^ "^ •'. 'I w in o ^' i 2 til c i s •g'3 1^ i« d d «' 2 (9 <1 <1 2-S e 1 q o r-; q ■^ ^ w m* + 1 + 1 ■^qqw -- N -N in + 1 + 1 ll V ■H; © o M fh' m r~-' ui t CO ooo© ©'lovd-- Air temperature in us in in >c«eoao en 1^ in^win a i <:moQ °i ■3S ai S V o 0 u o z E- < Q a z ij o OS a, (» C d o Q -<: o Cd O z -<; X u H S o o H z o § z e: o Q ft •.J c '3 5^ 1' s in?^' mm u J- »in ft 13 11 li 8 1 *3 C5 csr- 5g a s — 'c' in*# £ 1 C s o s < IE -Hin J 52 as c oin + 1 < * + 1 S3 -a c r-m or-; §3 1 r-in S 3 o mm ^1 dI to mm II 1- «-* a s - a O 500 m 540 ELod ot Ta»tirtq period n DURATION OF TEST IN DAYS FIGURE 95. COMPARATIVK TKENOS IN BODY WEK-IIT AMONG TWO GROl PS OF ADIT.T GROISE DURING PERIOD OF FASTING AND FOLLOWING SUBSEQUENT INGESTION OF FOOD The consistent gain of certain birds was due chiefly to a larger daily food and water intake. Such birds would recover their nitrogen balance quicker after fasting than ones which ale less food. Furthermore, the more protein eaten the more rapid the replacement of PHYSIOLOGICAL STUDIES OF THE RUFFED GROUSE 773 body tissue which had been destroyed as a result of fasting. The importance of the water relationship in maintenance of body weight has been presented in a preceding experiment. An interesting observation is that even among birds showing a steady net gain some weight was lost each night when no food was eaten. In table 118 are listed the calculated amounts (in calories) of the foods used which should have satisfied the physiological needs of the birds studied under the environmental conditions specified. The caloric intake determined at 6.5°F. and at 85°F. should maintain body weight within the limits of normal 24-hour fluctuations; and at 65°F. should maintain (1) an average body temperature of 107.3°F. plus or minus the limits marked by diurnal range; (2) an average respiration rate of about 63 per minute; and (3) an average heart rate between 300 and 38.5 beats per minute or an average rate of 342.6 beats per minute. T\HL1': 118. CALCULATED DAILY CALORIC INTAKE OF FOOD NEEDED BY ADULT GROUSE TO MAINTAIN WEIGHT AND VIGOR* Average daily food and water consumption per bird (grams) Daily caloric intake per bird Air tonuMTaturc Water Mash Grain Apple Alfalfa 65° F. 80-85° 22.6 26.0 10.6 8.8 12.7 7.9 29.5 20.7 0.9.1 1.08 90.8 64.8 ♦Applies fully to the birds iiserl in these studies and the eoiiditioiis under which exiieriment-s were eondiict«'d. According to the data, approximately 3.6 calories (biological value) are required to pro- duce one gram of body substance containing 0.40 calories at 6.5°F. Although the data are slightly inadequate for predicting the caloric value per gram substance relationship at higher air temperatures, it may be estimated with a fair degree of accuracy that about 2.0 calories are required lo produce one gram of body substance cdntaining 0.50 calories at 8.5'F. AVAILABLE INSECT FOOD FOR GROUSE CHICKS* Grouse chicks, for the first few weeks of their existence, are largely dependent upon insects and closely related forms of animal life as a source of food. Since their welfare is thus con- cerned with the availability and abundance of this kind of food, population studies of insects and their allies were undertaken. To this end samplings were carried out in each of the various types of cover that make up grouse habitat and under varying weather conditions from June 2 to July 1, 1936 on the Connecticut Hill area and from June 9 to July 14, 1936 on the Adirondack study area. These were repeated on the former area from June 1 to July 1, 1937. TABLE 119. NIIMBKRS OF INSECTS AND ALLIED GROUPS, BY ORDERS. COLLEr.TED FROM ,$-F()OT AND 10-FOOT PLOTS ON THE CONNECTICrT HILL \REA BETWEEN JUNE 2 .H LY I, WM> \ND JI NE I JULY L I<).i7; IN VTCINITV OF THE ADHJONDACK AREV BETWEEN JUNE 9— JULY It, 1936 Order Insects Diptera Ilynicnoptera Ilomoptcra Ilrnii|»tfra Colrcpl.-ra l.n>i'li.pliTa ()rll]i.|>l"rn INf'uroplera Plccoptera Oiionata lOplieniproptera Tliysatioptcra Mccaplera Corrodentia (*olli?nibo)a. Trichoptera I iisncl Kgjrs * Insect allies Araneida I'lialaa^ida Acarina Isopoda Clliloi>oda Diplopoda Pseudoscorpionida Total Area Adirondack 1936 U)-foA4 138 106 17.5 36 ■lO 0 1 0 0 0 1 4 8 0 0 71 29 SO 46 3 1.821 Connecticut Hill Sweeping 10-foot square plots 2.234 628 991 481 31.i 136 19 12 6 1 70 5 12 8 0 0 475 5,383 Intensive 3-frM)t S4|unrc plots 528 791 166 153 175 62 11 0 0 0 17 0 11 695 0 0 319 68 58 38 24 15 13 Sweeping lO-foot square plots 3.697 660 2.097 757 397 203 156 18 14 0 1 42 7 34 19 3 14 564 28 36 0 0 0 0 8,747 Intensive 3-f St u w 67 52 Z7 ■15 A 170 156 112 134 67 75 71 51 • .■!0 107 104 103 r.» III '»2 56 36 38 56 • 23 15 J 44 *Not Bamplod on th« Coonecticut Hill area. dNot aampliMl on Llio Adirondack area. FOREST STAND IMPROVEMENT WORK IN WOODLANDS IN WHICH A COMBINATION OF FOREST AND GAME CROPS IS DESIRED By Gardiner Bump Whenever the multiple purpose of producing sizeable crops of timber and of game are to be served within the same wooded area, it is inevitable that many specialized forest stand im- provement practices, usually considered desirable by foresters or game technicians, must be markedly modified. The first decision still remains the determination of the crops to be en- couraged. The forester may be interested in producing posts, ties, cordwood, pulpwood or saw logs — the wildlife technician, remembering his choice is limited to woodlol and forest game, may think of grouse, squirrels, rabbits, varying hares, deer and bear. The control of woodland plant succession to encourage the production of one or more of the forest and game crops mentioned above is little understood. Foresters aim to improve stand composition, quality, density and origin by carrying on cultural practices, particularly weedings, improvement cuttings and thinnings. Wildlife technicians seek to so manipulatt- composition, quality, density, origin and arrangement as to produce adequate food and cover relationships for the species which it is desired to encourage. The addition of arrangement is important. Recognition of the character of the various woodlands must precede ])ropcr management. The principal components are: crop trees, wolf trees, weeds, trainers, whips and under- growth. Classification of forest individuals or species into these groups depends largely on use. A massive, though decrepit, beech may represent a wolf tree to the forester and a crop tree to the wildlife technician by providing both mast and den hollows (food and shelter). White ash may be a weed or a crop tree depending upon the use to be made of it. Kim is seldom the latter. The important criterion is use. Determination oj Use The principal use to which a given area of woodland is to be put is usually determined when it is acquired. Recently the principle of "multiple use" of land under either public or pri- vate ownership has been much discussed. Over large areas devoted primarily to forest or to game production there exist, ofttimes, excellent opportunities to encourage a subsidiary or secondary crop of either forest products or game which should not be overlooked. In the past it has been considered satisfactory to attempt developments which encourage such a subsidiary crop on the same area and in conjunction with those designed to further the pri- mary use. Carried to extremes, this has, however, led to considerable confusion and conflict of development practices. It is therefore suggested that, as a result of type reconnaissance, it should be possible to divide an area up into primary and secondary use units. The primary * Originally issued in mimeographed form as Circular No. 3 by the Bureau of Game of Ihe New York State Conservation Depart- ment as a guide for Forest Stand Improvement Work on State Game Management Areas. 778 FOREST STAND IMPROVEMENT WORK IN WOODLANDS use unit might normally include the majority of the total area and encompass all save those individual spots which because of cover, topography, composition or location are particu- larly adajttable to secondary use crops. The following suggestions are offered by way of defining more clearly the characteristics of each unit: Primarily forest units 1. Plantations 2. Stands of relatively pure hardwood or conifers where game food and/or shelter is mark- edly deficient. 3. Extensive areas of mixed hardwoods and conifers where the crop trees are approach- ing marketability and are of the more valuable species. Primarily game units 1. Uneven-aged stands of mixed hardwoods and coniferous species attractive to game and possessing an abundance of "edges" or brushy openings. 2. Overgrown areas which have seeded in mainly to a combination of food- and of shelter- producing species such as apple and pine. 3. Old orchards, fencerows, and road borders, where substantial stands of food species, attractive to game, exist. 4. Stands of mixed hardwoods and conifers where such border on areas of practically pure hardwoods. 5. Narrow stream bottoms and banks where the species are particular!) \alualilt' to wild- life. 6. Small slashings or burns occurring in forest stands in which other openings and desir- able underbrush and ground cover are deficient. Depending upon the local market for forest products, the game species to be encouraged and individual site cover conditions, additions or modification of the above are. of course, in order. Once the |)riinarv units have been delimited, emphasis in development should be placed on each depending on whether the primary use is for forestry or for fish and game. In this con- nection the following suggestions may serve as a guide. Order of Forvsl Slaiid lin /irorcniciil Operations A prioritv of forest stand iinproxcnienl operations is suggested as follows: A. In units where llic (Idininant use is to produce game 1. In stands (icIicicMt in game shelter, increase the latter by: a. Liberaliiiri of supjuf-scd cuiiifers b. Creation of small slashings 2. In stands dcncii'nt in foods, increase the latter by: a. CreatiiiM uf small slashings, especiallv near coniferous clumps b. Liberation of fooil-pinducing tree and shrub species FOREST STAND IMPROVEMENT WORK IN WOODLANDS 779 3. Elimination of trees that are diseased, of poor form or of species of little value for game or forestry wherever they are retarding the development of more de- sirable species 4. Harvesting merchantable timber B. In units in which the dominant use is to produce forest products 1. Improvement of composition by weeding in young stands, both natural and planted 2. Thinning in even-aged stands of pole size 3. Elimination of trees that are diseased, of poor form or of species having little value for timber or wildlife 4. Harvesting merchantable timber Cutting Policy A. As regards composition 1. On any given site, in accordance with the relative importance of the two major objectives, favor the following species: a. For forest products 1. White pine 7. Hard maple 2. Red pine 8. Black cherry 3. Hemlock 9. Hickory 4. Spruce 10. Black locust 5. Oaks 11. Basswood 6. White ash 12. Beech b. For game shelter 1. Hemlock 5. White Cedar 2. Sjjruce 6. Mountain laurel 3. Pine 7. Yew 4. Balsam 8. Common Juniper 9. Den trees — beech, maple, basswood. etc. c. For game food 1. Fruit tree species such as apple and pear 2. Mast producers such as beech, hazelnuts and oak 3. Nut producers, such as hickory, walnut and butternut 4. Pulpy fruit producers such as cherry, mountain ash. shadbush. hawthorne, viburnum, dogwood, mulberry. e! = < ~^ = . * a. o^J iig ™ « o - a; i £ » s n ■' fell I- oS ° S o "•~.'a-a 1. J= y « "^ c3 >• , 1*^ in C_— o ' 3 O _^ Ml o ir ^J ^"H C i-= 3-^ •J2— 0.2 £: = - li ■- s = u = u £ C M > C3 3 OX O '- Ox*:; U Oj c e i: = 5 « i = .a^-n * =• £ ^ = = I. _ m « =— -n s^C-^e; ~ . z = < £ U -o ■< ^ ■r. O i >- *2 ^ a, c 0 M 0 S ■S X C era u .III O " k S W > o u b O W Oi H o o fO ° o o CJ — '-' ■EiS g£ o «IMIM M-^0«MOH' — h- o • r-H t-iM •M -o r- r - o> -* ^o-t o . £■3 «^ © . S?! r- lO-t^C ,— lO r-M s§ ^COCI [-, X O' ec O ifl r-io r- Z 0 < ccOC LijSu,iIcs ■ — ■ — Ho _, V c CO "B s ■0 0 ■^ ^x. m c tc ii« __ < >> a > 1 0 CD V3^ a H H o 0 cs C« ^ n K o o O UJ X > ^ O ,. O X UJ UJ O UJ Ql >- o S1S3N dO lN30d3d 784 APPENDIX o cc o tc M Q Z » o z HH Q Z 0-. 3 f— < O o S 0^ en i ^ H o -< >-l C/J H W rf) c: y. H Q z JH 1 a (S ^ w y n uo z M rr. Uri U z c >< r/5 H < en rr P L. O ■r, ^^ en Z 3 - O z o s 2j= 1|fi ■sis 111 z *: 5, *.=--^ r; ^l 7-5 -J ~ ■:'^ xj = ^ ~ — V 5-'=-^ c-" t 0— 3" = — 11 = = = 5 0 s s • Z ,1-2 5 ■axis .lis: J &5 ja to 5 c ^3 a c 2 o t-XS-XSt- I-IJ7 1 il £-0 0 0 0 000 CJOO 000 oooeoc d d 0 d 0 © ooooco II l- -JO* CI M ^ ^ 0 CO 0 0 r- >o c\ CI II It Number of nests OS vo ^ •-; so c> S\ « CO Cv 1^ 10 o 0 I 2 CI .1 II : S 1! S : S .-5 S : d d =■ II M C r^ rj CI X -? O' 0 so CI 0 ^ CT. ira so 0 c^' -: d II ?j ^ ^' S ■3§ II 0 a tf3 > : : : : : s : : : S =; : S ^ 1 S .= .9 ^ ^- 5.5 1^1 .S *'r <£ = il B< o S3 Z o c3 Is CO* r- ci -- X rt o^ I II Z O •< £U2 u: ::: u. u. ;r 1^ ^-^ 1 i r 10 o 3-0 o ^ - SiS3N JO iN30a3d APPENDIX 785 «■ -2 = = It: 3 = = CO I O DC o Q b O O y z u M « w H S3 Ms ° = = 7 "T^ ■ _ = -o ! - ^ O M C : I S£ = - ^■ — ~ -./: : ^ =■« a' • * ^ = 5 O O m ^ S — O o • I |i I ■ O c < 0 .= ■5 is £ ss J R £ f ^ >) *h 2 U3 •^ - H T 2 % 5" c 'c S K S) s =-a a r-FHVO-fc-ic:.£Mr:c^ ^ w m" C -• r: = © c c c c O* l^ » © X C-l i^ vc c^ ee «■ o E"=| 3V t' L* C^l ■-• •* z n •"• 111 •3 i S < r- X ? ■io 3 »- 3 i « 4: 1=. * III- 3 1 c ^^-r c L. ;;- 0 t ^. ■I 1 «3 a;ii.'H.'=.*2-- = = ^ z. - = ;^ 2 w Z 1 786 APPENDIX cs w > o o W 2 S5 sfe o u ^ a: Hz fcl w z " w c« Cu -^ 9 -J c« c« W O Zw H W r« a. 3 a o w CCQ o _ o QH Sz Uo 3E fes y. C '^ z o i. C- ^ ~ ■a j'S r- eecii/j iraei -*m t-~ M o es r-Mc ce oio n^ — H z = — < '^ ^ ss zg ■Si M^Crt r-eC-*-^f 1 rH r- M o CO 1^ o ooo cooo© o CO O 1 © u2 CO o o o eoo 3 — 0 r^ II MOSM OrtCO-^?^-- r--*> o C) S2 — t-Cl r-- X M X s n £. 2o 1! M oin f- ■-•«^0 00r-0 ift e-i CI Cl &a u^ >0 ?C -^ 38 Ct t-c r^ fn ■« a l-o n 0 V Q 1 ts = »« « ©3* CI t- ■- C) CI c — irtirt 2 3 Z o S3 . •f eer^r^ cit-»-*r^>': r-o ^ cirt CI si<« ic lO r - ^ -O sD O tfl 1- SCO O •J 9i o .s5 ■2S 1- o M C C t- M © CO Cl-^ O CI a r- -* iri tn >o ro ^rt 5w Cl-^ Z s II M MCIO -t i« * X S lO cow o« CO f 2 M ^rt- CIMTI — rcci i II -^ '«&■ M M CO ff« X *f CI Ct^ ^ ^ i-o I V 5 ■< sue UUEbSuZ N--^ Lm 5 ■ — . — ■ ' ' ' ^ •^ = « 5 cs ' ; 5 i- I C -2 s 11 £ a 1 1 ea i 33 0 1 H o o ? !/i li ' 0 * CI 1 = 1.6827 0.5602 0.0501 •5S CO s 1 c '2 V li ] 1| ir; (Ti — CI n — CI II Total Slope Cover type Error kT.s J 0 £*S J|l.l s « » a = i = . 3 = o| — i— i i I * ? i2 ■« r '• = : S1S3N JO iN3Da3d APPENDIX 787 s5 r- in-#'.D f- CO C'Unin CI --10 T- W o E* t- H 2c r,s — nic U7CIC" -t t- O LT r- O CI 2 ^ 2*0 in o in CI c ©OS oo o o o o oo CO o o o coc oo o o S3 =£•2 A 3 ,, -t-a — ■C X 1- ^^ CO SS 2 0 *j « PP c-i c i-r^ sec- =: wi-« r-ci o NC in C-J'* — Li Cl ■£ Cl •* =0 o — t- t' l-?. CO ■^eci-t CI WMCIWCI sncn C4 CI C<9 b 'f) ■oe- f^ " K " C *- "1 — ^^ _ ^^ CI X M 3^ CI 2 = ?l CO t^fic c — — CI in — l-O CI CI tinco t- i-irjxinec >CiC eo M «<- — rt^ w cici-HCjMec — N CI CI o 1 e. o s Si5 a c CI CI CI o — ■ irj — ■ Ci'; o — CI 2 0 H V 03 O-^M sC t.-: ?■ re r^ — — CI e- M r-* 05— -n — I->C OO'+t- tno in m l-n ■^ "■"^ — — "^ ^^ S X Si ES M rt* lO ( - 3* irt r; f re •n n in £ o Si h- — — 50 ^l-O iriMin t- — C\ o* t- l--r rt - X — le ^ - *CI I- r- -=•- CICl — — CI — — "" :3 UJ si ES X » 3^ Z C = 1 W ^I-u^ h-s3-'inc4t- CI«C o ii C c- ^-'^^a^ ^ ?^ cin c- c — CI \C « I't- c-i — — 1~: — — ■" 0 2 feiS SS I- TICI — ) Z o n ^ |=UQ S ffi _ " <7 C 1- n £0 Q 5=? c c ■i 3 ,o > ^ ^ X 2 o _I u. o o UJ a. to < v^^^^>}?>y/}y>}y/^>'VYm. Ez: w»^^^^^wi £±: 222SZS T*"*? lywvwiyw^ww^^vw "1 ^'''']''''ivpjj^ cz g 1 1, ^^^^; ' ' ' fTT»>TTT rr^;W('^/;'//r^;^;^^ ^^^TTTT^ T^^^^^^^TTT vzzzzz > o o o Q. >- ■J SiS3N JO iN30y3d 788 APPENDIX s CO O H O ft! a< CO o H »?; o ^ H CCi Z hH CC w ^ > T3 o § o J Ci. '5 C 5 W^ 0, ^ >^ CO H 2 >■ S eaS ° T 1^ Sh t3 CO E w z£ -H ^z ^ (W PIQ z H tn P o tf o o u (IH K O E ==i Z o ■< H U ce a, cs u Z. s >.ty — .s •^n. 2| II fc-i S-2 *.l *s^^. ' i « E ° "5 ^ S o c Z *i m a; J Z r- i .2 S CI 1^ 1 = o.2i:(:) 0.8816 0.0492 > "SI 12.7166 0.8534 9.6976 2.1656 'o 1 ■< la W f -H ■» =1 0 a »3 > 1 5 z M « — o = l^m 3 = - = 1-1 s = s - = ■■= s 0~; — ^ X O z o H o o J APPENDIX 789 •a 0 gy OJ S'c 0 ^ O 0 p 4) •^^q 13 U o2£ 5^ «-o = i| 5- i5=^ S z o 11 5^ — JJ < ^'^t .2 o Id tn'-^ ^2 a. £ 1 = H Sa a— ^ Z = ? b u *> ^as Cj3 ** r.-= V £-^,'K -0 2 Oj oi ^- 0 S-^ 2^ S'5 1-2 -■ ■5 " z r. - -* < s - s -3 C ?3 y ^,.2 t- ■3 ■o g CO Si ■*• '■J ■" fe £ * =5 -* S 3 .«,S^ Oxi-j-:i ^ CI M o o s s o t o IP 9 -* O lO C; CO « -; - t- CO irt M ^' ci o o c c B,«-= 1s| oooooo^-t^c -f o r- c) CI t -D ITS C-) — -HO^COeOTJ•s0 1-lOC IT loS w t . --"r-iNiN^^or-^h- a-oiE z J- u a s ? "S o <-CiOOOOO=:0 0 — Cirt-tLOOOC UI 1 o CO •< H JS2222222 > 2 5 f2 ■< ^ c M •* lr^ o o PI II a* 82 :5i .2 * I- -1-2 — C 61 ass a £ > a S ' E i ^ I UJ O UJ o ir u> UJ U- TT o o o 'J- 5: 0 or u. o l<) UJ 0 z < 1- o (S) OJ Q g 8 § 8 ? lOOd d3d S1S3N lN30a3d 790 APPENDIX S trSz ■15 r. z i c z o c: fc. 2.^ = - ?=■£>, : _ ;; : ^ >>■- — ^3 3?.= '0.= t r _■— — -. 5 . > ^^ L 3 V; = >TI £i £ 3: 5 ^ t'« 1*^ ^. £■ 1 £ H I :5 = cc ^ c 1 '^ >■ - % -} ?-S!= 4 < UX'S < a s " '- i' -1 w S = < >. J3 S^5 ^ :mi 70 >. < — ^c = aj -a if o H Z o I W O b: o b. b. Z o H •< O s l|1 62.2 20.6 11.4 t.7 1.1 1 sal lis l"3| 1-.' iii^s^n CO Number of nests ,- « ^ t< m «■! CI 4- 1^ ez m is'a mil ? r- ^ 2 c^ ■ ' — to 22S5S coco© * i IS So 0 = ■at V o 5i a«2l iS o -., *• £2^ — ir" 3^ 7. A' E ^■5 = 133J 01 y3d SiS3N iN30a3d APPENDIX 791 "3 0 t- W#vO h- 03 ej in »n ej rt if I ^COM i-co L- S^ l£lf "t CO-Ht- OS 3 o "■ CC ^ ^1 r-— •* W I-- M Cl O ■* * CI r^-- oe-in-*t-ec a- o M^ in « o 0 >?♦ S . H X C-l in wco o «n o o o; \c en en 2JS-S •* cio>'-^ r-" d in d m ■*' d d 0, C-- S « OS 0 '^ f-l --miN \0 q c; ac CT; -t in in in d ddd M ci d d d d d d f 6 i « «.-<{?' -* CI in o r- ift Ov CI r-* CI — OOM«CO-H r- ea rt rt p *0 d >>» J 5 I u X u i C 4) Si ""o .= = a 0. 0-- f>\ O CO -« fl -C O rt \C c q <^ •4 u-)— *t-.' r^' ci in -* iri (^ d OS CI — -. 1 € & « S s Ov o " ^ C-IvOm CO^O'^e^A^O .1 ^ B OS d d<6<6 e^i ci d d d d d d 5 i S 3 > to a ^ CI sC r^ M -f C ^C -O ■* ^ ^ a CIO Cs OS £ o a c >•* 5- a c o ceo ddd omocnr-o t- —■ d ■* — r-' o 00* © CO 1 a. CT- Q V 2 s t-i o ^ o coo ^c^l qse-t'O •* ■* g d i6<6<6 dcioood t^ r-^ in 3 o c c c X — o cnirirt r' r- « M Cm 0 0 >•* Sill M inoco \C -- O op CI vO 0^ OS CO m" d ^' uiwd'^in—' 09 09 VO i-i -H W .-1 t 0-5 'V ^ IS (-. (N inOM CI m N CI M o CI CI d d><6<6 « ci -^' d — ' — ■ OC CO c 0 '^ .§ 1 = pri (MO-- inoin--in-* ■* •^ ^ m en 0 a >i* S 1 " X t 3 S M O •# W ci cc q ^e so CO q q «' ideisa r - ci d 1- d d ci ci CO r- I/) ■rf in in-* wo ■o so a, o--= 3 - OOO qqqqqq d d en CI d o q CO q CO cy. f-»rH M m in o 'o ^ 1 C-J C -*■-! o orncjoo CO 09 — — CI in in *o a "1 -< fflUQ h ■1 U ■T3 ■g TT ■r 2 ca B o "5 a CD 0 5 *^ £ >> '1 a E? ■o 8 3: ^ 3 00 0 H ^ > C« 0 H O o Z o5 N — ™a B.-0 ES ■9 » 2^ o •5 o » ^ a n.S « M ^. M - >< 0 t i; ^ .J o <■ • = 3 S II Z < i: -.2 .J Z^ s - .2 — - -i <; ~ "r c -J u ■s^ 3'i '£ ■^■>. .. = < •- >> s = "c ^ = 2 ■n ^"5 ■ - "V "c s is ■=c ^IS M* ^-= ll ;; K ;? c i:: >> 5i-r. Z£ r- = r- ; 0 f C] c c -^1 so -* -. -^ r- r- S S 8 d d d 0 CJ II 09 -• tn — ■O -t CO '* sO "J- CO m CI tn in -: r-: w' CI — ■ "3 = Cs OS ■* so \ J? u = 1 1 1 .- > OS pi Si5 en 30 •* ' a c :■« = .^ - UJ LlJ X < g i33d 01 H3d S1S3N lN30a3d 792 APPENDIX o z O H Z O •J Ed DC O Z vo I— CO Z o; lei Cl. ^ — 5: umb ' nes per c- --rt^o 1-1 — p- M M s^ doM !>-<'© — a ■*' M ir O- M ec O S CO Z o - cs cn |3 c i- H X cc C-f — h-C'I C o c Ov d—' — ddc d 09 c- ti — c 1 t 0-.= S5 ♦■ Numb of nes per 10 fee ee CCl-* ■*£ ^ 1 a. =■•= O ooo oo c o e o ? ? d-*— ' ^de iri g si o zo - ^^ ^ ta ■Q 1!! ~ c — — £ c = in r- e t- X ON 3^ Z^ i ^1 -< XUQ r » M b . c ■| c " i^ c =1 0 c a j 1 > = E ■oat - c i 3 a . 5 s2 H Q s r.- >- z II Site c . fifiO ■o a ft li « = 52 1 = ■i'3 8.S 5 = S « S-fc APPENDIX 793 o H Z O l-H H -- ^■2 W c «'^ ?r P, p^ z <^ Z o aTi Oh S ^■i O H -< W H Oj CAJ Hg «g D W Sc^ '^y W " s z J w fc a, z o -S ^o ^E ,, a ^ o Si ^ Q Q w fa fa 3 Bi fa o z o HI H < O o ^ 2 M U be r" o ~3 -a ^ a; 3 £2o c-5 >i CO bL b -2 2 = SS t = = i, '■- ■ ■ ■£ a Sf ° S ; =" f.2 «i ' £ -* 5 i C c ■ ' at 3^ - i;^ n^rS «2 -^t^ limb \ II »i :! £^2 S ii 0.1792 1.6026 0.1163 W 5 ?^ r- o nS 5 U * " J^' — — ri II ^ "■ '• T> ■Si II 1 j2 s Q _s S 0 "o l 1 i. g 9»- = o § S § 8 133J 01 aad S1S3N lN30d3d 794 APPENDIX Z o o Cd fC >* pa Q o z b z Q Z O aa o z >o M O o^ =^z O z 5 o O O <: CD H fS o 1 .! • 1 ■» c CI *? o&>c^ 1- ^: CI C^O ifl w Eg l"3 ^ r: C ^ !-•* S "" s |S| C C ti^irt ej u- r^ C !-: 5- ^-0^ « o u S « s i .-.':.- O-f 1 "3 ill ^ -*t- c L^sr^ziu-: — tC r- ci in 1 z-sJ tea -ai cc = ---^ — r^ n i^ ^•3 111 ■t- C =: t- C ?■ C O t- C c^ OO t- Ov o fe-U c o s Cir::^ oo » ?3 ^•sl - a^~ •-^w>CCirt CI-* i _ teS^H (. o |!1 C — i-^O X s=^^?lCl — — C) -* ^ in tea ss c crto CIKf-^.':S^^- l^rt vC ^•i2 S'' S r: oi--* g 111 ^^ 5C C-I O O ■^ MiO 1 2 ill " •5 Z oj: ■< II -* i:t\r.\r 1-^13= cr-: c ^sO in M in ^ s Z'3 eg >» _■ < £UQ ub:u.h.;:;x ■■"^ i i-S •o 1 o _ "S i M 1 1 o o U a 2 <; 1! CO ^ 2 '^ o"3 ■02 c3 a u at— '- * o s <2 :2 £ 3 t- * = t P s; - •^ c; u = S = 3i^ 8^» 0 0 s C aiHOlVH S1S3N JO lN30a3d APPENDIX 795 o 3 w M Q Ed O Z b Z t— I CO -s! Q Z O ^^ E c Z a: CO M Ov > O o CJ o-. z 1 ^ ti o H rr: ■^. U CO o a 1— 1 H W U a f- o H O z H- r 0 s. o u u -c CO H ro w z m m » O ti O b o u H < lb ■3S 1 = § : O d d 0.0277 a i c-i ci CI C-I - d d i-t ^ d d 2 6 d cr-a w o 11 — SO a^ '- c-i — vC i/^ X in H^ ?* Pt 1- — C X sC -*■ ?^ cs •*' •* d u: 1-: ^ w -f o r- — rt ci irt X t- irt « lO O ■^ « r-; M 1^ ON -^ 'f d ^ w — L'; r- o- -* CI C o o — § = S S fJ a<' -* rt d •* r- f- in 11 lO Cl — CI — ?n — CI Cl « M c 2 c a D o e2 3 'i > > 0 'Sc U t in tl ^ 5 c d S r= d > II u '* > >■ li o p^ o d CI o d II 3S CJg 5) M S3 s ^ ^• -2= to.-. "o - ^ ex ^^ ^ MX -. C O n ■"■=ii Si:'" »_ » « Be ■*• «r ^ ♦^ O"^ in 05 .2 5 =>'-5 S H .a »;:-5 „ a 2 "J.S ■» S S Jf js-asf J hS8e- §0 wz c-nH c- ire; w z z-o s z « 5 z.C a; t t; = I z ?': &3 " £ a, . z = c3t_ ^ e.:=-a « £ £ o-- '-•^ S r ; si-' i..z*S=" H5iSSS"o ■ 0 r. ^1 796 APPENDIX i3 2 f2 |i "^ •^'* X s Z 0 •c §5 11 irjci — — — ci X X « Z = = 1' ess o r-c o \CCJ — ?C*?CI -o — M ^ CIS ^inw * vn ift sO >c lo U3 "j? B. oj: ESs z-oi 2 eg ct — » M S3 Ec a. 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C..2 ^ ^ — f « ::^'S ill 1^ iij Z a t^ ^i 1 2 ^ r-(y I- i-mmciTir-: ••M r- CO in s| -* oin 0 i-c-i-*?^ c c CCI 0 \n H Z-o , b o> li ZS t) 0 Z'o C p •-I f-«ei «CIO^ CI S-- irjco 0 M p^rc r- K COfflTI-*-? ■^ 5 «1 M «« w 1-I?0-H CI — "t Z-o **'€ - V X •-( r-^o CT' — «MO- "*■ VOVO 0 0 CI ^A -0 B „ ^•3 •H 8 Cfl ■0 11 is ■^ccci |, fC — -■ — f^ S s z-s M «inM — -- Ov CI CI » tot^ 0 0 X =^•0 3 0 il M t^irt 0 rf CO CO — 0 h- «^ e\ w „^ M M« r- ^ » ^■3 II ^ COlOt- 0 'C W lO ifl M MCI 0 " 1 •f OCOI^ 0 ■* « -H FH NO 0^0^ 0 c M 0 * ^•s C P y « t-CJlrt ■"■ "■ « Z'S a4 •-• taiH -< fflOQ csjll:[x.^cs — ^ -0 iH Ci] V t- CO c a 2 s^ CQ c 1 ^ m .a 3 0? 0 H H 0 0 s [« 2 . — at s C - „ o X 3 S - = S •2-= 9 J S §.£ £ V 5 ^ E soooyg jo iN3Dy3d 806 APPENDIX s S.2 s i£ = 5 |-5= = 3 iP^i' H z Z O Q Z o o a: a a o H •«; o a: Ed Q Z CO Q O O en O 3 #'•5 Q-e f S 3 I L. 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Jt ^' ^' '4^-55'i I- i*:^ >i j; >i — r, .^ • ,^ J. 9^ ^ r? Ti — — « s* Irt =•34 CJ X 0 *^. £ - ^ ^ u as = ' C z -^ ti X s 0' > i J « 3" s: X ?l S' X 0 c 0 ■■' " ■ J - « " & j?^'^ « •:% Tl - CI CI i = .t "i ^1 ' - j 0 k 1 -s 1 U. s*s| \n t-: •? PS r- f- ! 0 0 z ■* _ 5 5"- i nx ? £ r X 0 » E = S \f: 3" CI ^ i ^ 1 i >: js r" > I'si — ^ \'i o* ae 0 1 ' > r- • ' 8 ;? i - "■ " 1 g rs « u 9<<- 5 ■c ■c X •c c r- £08 X 1- 1- !^ 1.1 i ? .- « s 3 c fi w W 1 Z ^ H ' o-' 1-! t -= 0 ^ ; i'i ^^^r^^^Fwv^w^^^F^^^^^^^^rv^w ik*******A*«MbM«b**rtiMMMi*********A*A*** Z o UJ O :i--J cri- uo >< OH OZ i8 crO L.I1J UJ cqO 2i 8 ? S saooaa jo iN3oa3d O O o cvj — '-' APPENDIX 819 CO i e2 Qi =3 S ■o — MO d ifi d COM C COC O =0 CI r- CI n X eo r-; 0\ -^ 1^' d CO CI — t- ind o © i h-inso ci ci (-X-tO-?X vOOv § m ci « in 1 Lit X d CO — C) ©; d t- d d ro JO d d JON — X dd i : 6 d irtin-* odd co'fj'MXin — do odd— enf dd X in ! 0.= ZS O— M — CI — !* XCI XOff- c r - CI o o M ny: — d r-'d i : d : o Z S I. (M C d O oir; ddd Off- O Oa-ffN X C) M -*• O -* d d d d — d m-f d — r- b-r*co -Cl- oct — — *?5 o ci 2 CI 1 Hi sit d r- o o u-'ift'd r- X o — X CI ■^ Cl ©•_ O X X d X ci — lo d — — CI OCO en CI in'x i : d ; o .11 d x« — ocoin c> r- r- fo CI M — — ci ci — -J CIM d '. CI 0.C zs r--^ X — om X X CT- O CI S' in in CI rt — CI CICl m t- — CI sd X o H 8 > i 1 O c _2 c s 2 > aSi-ilcs c o V 1 0 2 1 1 3aDV d3d a3Hsnid 3Snoa9 AO lN3Da3d s .^^- ^^;» 820 APPENDIX o CO ON o H O b3 Z O y z o CO ■<: CO n Cd CO O A O n Q (d > O o O u a, i S s -o Z o ■ z = . 5^ -, 5 ■ * = u 1% ;d CD '"' ^ H - 1 0 ? - S; i .2 ~ ES §3 I- i' ^' Is j|l a! ^ S . 1 E *= E 2 iiS-ES^= ^ s _ < '■= z Z. ' < 5 • o C^ = E^ c I i-= 1* = 5i- J i 8 — • S-^ ■ =12 "S!!; Z o c : ^ ci r? -- - £ = E "= sT ti; ill « u a. * - a; 5. d — ci ^ - U i = ' — ^ ^ I i >>.— 5 — _>.; >» >. i^ 5 ^ >.*^ "^"^ = - ?—■ r S = ■- u,_= ^ - *,E :l-2 •<.= inC tfi-< -«:.S -<= loU APPENDIX 821 W ^ life CO -^^c» -# sO -t C< -H Irt 95 ^- © a CO *. ". "^p irt OS eo -* n ^ ^SD o § o LTJ N d eo" t-'Mciinci --"r-' © c* n © D ^, a = ■?£ _^ MCC-" M oseoscofo eo© S o -^ot •#— 'MCO-f OS r-T*; 9 > ^' d d Irt t- r-^ ci d d os' eo'-t d o 0 Z C= 0. " 1 Ill — Ovifl mi-ir; •*?) -o so so © r- CI sCO* ee-t coc> « so snoN © © cdcicd -f t- •* -H* d 33 end d u o O 0.= a L. s II i 1^1 \o so coo o-ooof -?■ so© o -J^ln in --©©«!» — in © ^ I- ■-■"dci ci-* dd 50 d -f ci d a. — CI © en ^ II i 1=1 Ov inw-* Nt-msOsO>0 •*« o 3 3 < M ift-Ot- ec M ei so OS M eoso o d « co" t- OS M os' in d ws dos' d o 3 II ! 8 = 6 o ©inc) m t-r-o-T--M incj © -t i-ir;« in o in £ o 1 - coos © d d o^ t- oi-rdt/id r-" os"d d ■^ o J3 o.= a a o s U II! 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S.: J3 t:_ pi _ - il t = 5» S 2^5 3 t^ S « 3 i'u S 3 ; a- = "si i B ^ i. - .3 « H ■< H S E- 'S z ■§& B-5 at'c i 7 9- i^ -3 a 3 9i *> SB C . -2; ** 3: s i =■ 111 I.- off- X = -^ 3 3 « >. -3 S 5 S. 3 =. - - i'"3 i-il i.-0 Is 5i o S: 3 :£ J3 3 ... ., P S - - ? - ^-3 ■ = 3 .i-sx a ■- .a .Si S'SJ -a i£ «5 ■:; S ; 0 » °S3ie£ U-3 - _ I- iT = _ C-r 3 «- -. T, = = = _ —• — ->>- yj 2'-"*C3 >, |.||"3 =1 3 = 3 |s< o^ .:? .E-- -3 3 5 »> ii ! =i ,^^ * J 5^ .2< . -5 i = = S"S-3 t=l^3, > 5= -3 M o6l^ < = rr. 3 "^X =£_t|_! — 3 i . * « X t- ac— *- =~ t i 3"^ i a 3 E_ 3_|i^ >. Ml a " = L- . 3 5 a aj 3 u— o-5~ S 0. _|- i £= fc = X .i 3 , picwr^ufflx.-"! i - *i -3 o ID o o lO o o c>J I Ul o (D 1^ m rt IT*!"* 122222: t o z o < O in o o O CVJ z o 2 3aov b3d a3Hsnnj asnoao do iN3oa3d APPENDIX 823 o CJ Ed Z Z o o U ^ Mr-o^ re M -t »C -t so ^ h^ ■c "n^^ 0 u MCirt en M u 2 = E H o OOMO '-e e-i © M in tn eeo o ^i ^ wtn sCinMi-— d cm o — CJ n 13 1 CC s^ ST? a-S >eocoo>>e-« f' in M E = c"0 n in oe-*« 'ccae^*© O-H o M in — ^o o< o* o in o in — M o •5<^ ki .— -H en o i:^ 5 ?3 S^TI «5 E-S CO oor- «N«r--#fn i-in in On ^ —•-N ci h- 2 = r? C 5X t-^d-* PI r-' M sD « « FS a.f= is 1^' S!!< E-S t-c-* — t-f-jo — m ON r- t- n rt o> » 2 = pi '«' ffiOQ •c ■a u 0} ^ •c sc 1 u g s 1 ca ■1 99 0 0 t- O O & xn 2 EI Vi'iyiVi'i'i'i'i'i'i'MyiVi V???VVV !■".■.*■ ' t ■ 'y^^^^fl*! >ij>>>i>>V>>>>ni'i AAAAAAAAAA^ < C, 2 CO (M (M ■ Ul o m o o o (M a3Hsnid asnoao dO iN30a3d 824 APPENDIX < a o a T3 to Sri S^r-.S = . * 0 ."r^ « 3 -o S c ■- 3 £■.= ■= >.- •=> |.s " ;s i-j HS zJ s-Ss=-|.5S= sISIfI - z=-= >: E I? ISIS i-.T.ii IS S 2 OS ■=■£ cS-'Ji^f ;i.-; '!£-5 £-5^ r, ;?. "^ ^* 5-?= iT =- -s : = ;.=■? hR :!: j4^i-lll:Ei£ii3i^ =:" = « J I ■' = = ■-"■•"-.. I hrf.?£f.'j = >-! CO |C« _ _ q8 ll5.= al=iJii51l=ll o o u. o td 0. APPENDIX 825 2 a * te-o S E-S ^ 3 3 z= « « 1 ■0 S-n T^S a-s 3 3 ^ z= i J, 3 ll t~ CT. — C- r- CO X c c c-i o^ cni- !-:« r^ ltmj- ijir^— m •—w I- s 3 3 2 = ^C « H S 5 o PI e M xc-ir^ f0r--*O3D — SOO^ o C_( r. Cl CJ C'i O 0.= "^ u z z s« S'S E-S CI m o» in — vO -C CI « in in 1 z= ■^ ■^ -«; c-O Q a n » c-io-o u- vC »■ se CI o i. tlft Ift e- — in -* C) M — en o ki c-ici — ■ td >. § >,=n 0 B E-g = 3 2 = CO — a^ inr- — o- -t^ en en u •^-f'^ en Ol ©i H F- Q H « s « weo-* o f- * ec a- vc — o o H » eici — o 5: {£ = ■^ u •c . CC S- S-o o — O"^ SCO t-o^ m-. m E^ «■) Mu- L^ i?• ce m Ln C) — fnm en Z = z 2 a. a! 3 -*■ eocof* CI — so C' eo o O'O o ^ .So' CICl — o en 5^ 0,= Z S < S'V >£ eoL';o^ -H m o o^ CI r- -f 0^ in in o E^ -H-tm^rncn o 3 S 2 = ds 6C U 3 in h-ON ee ec o* w * CI «o^ o M CI cue o c >j P C-G ■^ Q <: ^_ >H s- E-3 „ t ir: c: n 2 = Q u S >» 5 < ffiUD :i:u:b.blc = H " 5- o o ■D c ^ « CO T5 O 2 ■B c £ .=^ U3 g s M C c 1 .s s 3 « 1 K > 0 t- H H o O M 2 o to r///////<'//<'//////////////77;7^ t fcfcfcfcmt»»***>t*4 «*****»<* ^5^S X ,.f'4V*'»i "E [^////'^//////('/^('('/^//^//'^('i^ (■■I 1 1 1 if '■'i\'fcV*i'fc' re .^ '' EC 3 O a to ^ CTl cri O^ o 2 CM CM PJ CVJ PJ en — — rO CO Li. ° o oo oo ijj ro ro to to to 5 CD -8 Ll. O X uj 826 APPENDIX o H a u H 00 i, ■< m Ov o-r ZS S2 & 1 mJ, Z td Sac «-«: £fi J ■=! -3 O — fiS ;fB 5o 9g x^ paZ o QO [d • O O Eb o u & >• H ■iZ =^ =:5 £S r : if, =:; =;; i c ? < ■ Z ' < c « S = < ir, ?< ?< >.r r.'- ^3- ^O" — CI ^ f 1 51 1 =■' - s - ? ■?» ; ■' i ii ui si ' n — t,= "■ *« E^ -^^ r^ •* ■* Z = --a a n ■* MMOv t~'^t~CH^t^ »•« © CO- O r- Win •O in -* M « sD rt ^ c ki Ci"-" ^ c Sn >.':^ lY 0 3 3 Tl J i^ o u r- ocj r-i-c-j — — r^ — in 13> § 2 = H M CO c-ieew « M in ^ CI t- ino^ © o r-ot- CO w* — ^-e CI en © -0 . &,= s2 SiTI E^ CO sOff- r- r;f sC-*»n •* » © © ^ ^ « en 2 = II *« ^MSO OOvOCOW — rt« © .9*- o «■»*•« ooMn-N-or- CI c i^ 0.= '-' TI« fc"C s"^ V- •f ^-*-* Ct 5^ •? O" «cc«-t■!J• ■—r- ■■c o z= = ■0 c £ i •* in CO so fc — M-* © .4 in -*•?) so M CI in 0 3 o O O « 2 QBHsnid 3snoy9 dO lN30d3d 828 APPENDIX < Q o o H Q A g ■< ^ p I - W S^ OH §- QO <; w «o U m 3 6c "2 - = "= 0 & m o >> a. m P . - J= e 1*3 = 0 C I. 0 0-3 3 OX^-r 52-3 5 _• -:: •- ^ -^ S. « (0 St-* sfss :I- .'iSi-S II- =-1:1 — * >''s a ? 1 11^ * C.£ - sSlI «.=■=- : 5 t - ^ - jc ^" :.* = 5 5- ?i; £--=i 5j-ssEi * s. ; >■ 1= s ; S a t-^ 3 f3-3 S >"- i — -.2 U-^ 3 ^ *5 Z c ^ ^ * ^1 So -T-S • >. li a g s= ; * c _ 5 a ■ 1^-^ I 1 > e-3 > ^ 0 -S ? 1 '^ ?r = Sf ■ 0 =-j .3 ; >% C 3- J= ■=i'i*u _i 3-a i.o r ».= ^■5-- i =■ i H=" g^a i-- S a = ■3 -3^ !3 C c C" "■•■= •«- 0 — E t S2 33 1 s H < X X 1^ 2 <: 0 X z ""'o _; £ 0 <: N r- ■5<' |2 c^ ~ ?x S— ? "■■ ll < — ti s ? a 1-0 - s v » -£ =■* >>«— 'li 5 ■5 = U. 0 **_J '^ c § 1' •ESS ■*■ .-— = e.- > o o o u o CO On 5 u H O W o o I >- ■< a o m I— ( H O H Q W H CO CO o o Q <; IH PQ Q o o Cb o lO IH 3 H ? 2 .2" « si (- z .g«iE~- = = « 2— S > I. >. 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I N III. I ii ^ 0^ Q CJ 00 CO CO (M ^ 00000 l^ ro ro pO fO rO 5 tc cn — ~ ro L . . .•■■.. . ■• azsZk "'i'i'i'>'»'i'i'i"*'i'iV»' c^ kwm:;:;;:;^;:^^ ■n^;,.>,sw.„,;s:w;H k^/////<-//.v//<'»>* > o o UJ Q_ >- l- a3Hsmd 3snoy9 do iN3oa3d APPENDIX 831 o Q Z O U z o H ■< f- . K « c H ■ ^^•|l|| 5 = ■ ii" Z2 ^ b (- C tb f i 2 iJ us To « n -■" ^ » ■= - £ -3 J, « = t ^ d r i* s *i 2 2 i o S 't'« £.5.s:2$ e - * i * S St i', . ?£=:= 5 1 • bt o u y r"-5 ■se c - a = «:=.?= -5 — ;i — M z < o 03 s» s t^ a b PS w a. S • W vo ON 2 "" > w Q t) w o a Q • n Q u 1/3 a: w > o o o H >H H Qj-d CO \on » CI 'f o^ r- « ci<c Tf o ■* .o — in Cl'^ o ClffI — o 1 &,= & b. w OMn-a OM-i» WO^O^ e-s eo oo^o c*5 w cj ^ -^ r- sOC^ >o CI CO z= M M 1- l: -C ffiUQ 5= K _ ^ -u _B •^ 0 ~s 5 ta ^^ 1 w 0 ^ , , ex Ic J3 a H a. > o 0 h O O X ^ * ri; s > h8 aaHsnid asnoao jo iNBoaad 832 APPENDIX = - X u r^ : - i '■'T- Z o ' ? »— — .r ='": ; « i *:= . 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J * < Z < , < u. i-a SiT, 3 e-^ r - ff 1 - ox-* £ M« MnO •^ •-< in h 3 3 z= 1 I'S "J* 2g 3 3 — e* » 2 = 1* irt ^^ eo-*irt coff'^>e'-io — N 0 V 3 &= ■^ » c tf} c^ 3 3 CO C 2 = .2 c'O "■S C sCf- ^Oirt f^»o» •V 0 b >> e.= ^ u X S fe-o E-S n \0(mn r- < 3 3 "^ ■^ 11 m r^^c-? coei 0 — Ml■- Ne« 0 „^ lrts^«C >t o.<= -^ 3 u S^ B^ in cg*oiQ « — ci cj r- 0 ^C EO r- CINC^ ff>. CO — — ff^ 0 0 3 a Z = ■* ■* = ■3 H —1 C-J-iK ©■ •* 0— ?^ "^ --0 » irtMift eg -^ -# i/i — in Clio 0 NC^ —' >t Q.« '^ Q a 3 « SiTI §■3 «»ft M m ^f M m CI c ■-HPS in so ^= •"■"^ "^ NO ^O £- >» ;■ ■< a:uQ aau.SoK N-»-t H T3 ■0 s 0-3 ca [d cd g Bfi 3 Oi ^ ES H° 3 d m a. 3 0 1 h 0 0 & 55 2 — ~.| ^ 3^ — ^ X >* >^~C z_ >>- >>- llfllllll 7: : c •< : r. 0 y. s - C X O 3 H ? 000 dO lN30a3d 834 APPENDIX n >> u ^l « 'CMO' «©MvO — CT' 5 E-S CS=-cr^^ CIO »• "I 3 = ^ C"! CI rt z= " "? Jl ff> Min* ONCsct-r- CISC t~ CO 2 = CI in se in r: -t cim "1 % 'f « — W e-i c — ?c — w rti- o 2 - o rt WW t~ M r^ « ci 0> ^Cl o ss Ik ^(N C-l n 0 « S'S M iftino* — -f-s-emo O-Os •f m E-g c-j in in -- — — o ^— C-l r- r- 2 = % CO ^4 >4Kirf c- CI o* « -* in cm © 11 II -^-^ -}■ o si SI? a- -^.- >fi O CI CI Cl M n m vO 'X.— 3 = 2 = © o II M — c o -* © CI CI 3^ — c t- © icr^ sC ■c S| 2 u u C E-5 «.= 1— t •** — 1- r~ CO £ 3 = z= CI M C 11 ei Ov«« ae © CI * o -t ^■* o CI iCsC-* in « rt CI CI o cjrt © fiS CJM CI o ^ 0.= S-- S-5 u^ •f c^ en m m 3 3 Z = II eo OMM OTpcin* w t-o © WOD* ■* CI « -* CJ CJ «« © 1 M «MM S5KiS= mm « CO 'f mm — i^. *f 3 3 2 = ^1 r- l-P5"f >c CI r- o CO © r, C« © ss e.= Q ^_ M so^eci ^e "t » m t' — MO ^ cn ■«* »> infi 5 CI 3 3 « 2 = fi K S < eauO UUbtbU" ■— "^ 5 .5 ■V f4 "0 V a u: s 1^! s a ^ 2 7i 1 3 < X 1 1 eg 0 f2 H O O s X 2 • o If) o o to o (NJ s z o o o °- ^ .. o o cc K *~ LiJ =- yi to CO El 'I ■" Vi'y>V*V*'.i'.*'*'i*' ^SS^5^ o UJ > O O XQJ LJQ_ a3Hsn"id 3snoH9 dO iN30y3d APPENDIX 835 (n Z O H Q Z O O Q Z 3 ^ O ■g a:: 3 o C S O N^ £ -j! « > 0^ OS S a CO Q 1 z 1 3 < u Up: ■< O H« 2 .J o Q -i! H o >' w oa 2 Q z H o en o & a. w > o o 5 0. s p t - . ~ H — 1 ./ ■ - ■52 'J >• si ■? ■- ■= (- £ £ I ^ * 2i.= x| z -J* ''. iM-S is ■ 3.:; = .= ■=•: 3 S z it ■r. 3 Z /• = " ■ill-::i- r C = ^ i ~ " i JE-=i r, -2 C:^ = ? St = ^ li I?- ii |i El Si !■- 21 y N il i.-i =.-» s-3 4 -- =.- s « ""■.5 =~='.aa-c - &* I* ^^ V '?''-='? ? -^ '5 = ■?■- ■9- >.i - ^^5 Sv-^s C;. Ct 2o = = J •■=! '■= _ ^ ^ ^.j = ^ z u a — - ■«^ -SJ i5 I W ■* \fi I- 0, 836 APPENDIX aj-5£ 21 •-.: i-H ,iip -: it Zt Z-Sr^ e..z ifl^ 35; :='^^|!C-«=j::: .J < ■ < ■ 3 t « i £' 111 ^ u,^ ■ g 3 w 3 O J. OS o D z 3 O CC a cc o b Cd CO O c I < o » o z e u t, e c z c CO •< V CA ^ ii ^ r- 0 E-S 0 W M » H 3 3 « « \o « T S:-= .M E-S M z 1 3 3 « Z = =7. CO w 0 S-S « ^ 2 2 i= i's ** w E-S c CI in 3 3 M re ^ so Z = ■■ " = "§ 09 M 0 r- CI 0 g 0.C is N M n i z= I— W « CO CO 1 ll OS © CI 0 § ; u__ I'r X r- in -* 0^ a- M s-s CI r- c 0 3 3 CI CI z= 1' rt r- 0 S'S » 0 s a t= ^ ^1 CI s X s in X r^ 3 3 CI CI CI z<= •'•« c S CI ae 0 g-s 00 s &.<= <& II « ^ ^ » o^ :? xn gj; 1- M 3-2 ■" « r- X ■w u X ft 1 a 1 B p 1 en 1 1 ■» 0 i n •>■ u< Z. 1 4:h i-fife-,^ t- i a3HSn-|J 3SnOdD jo lN30d3d APPENDIX 837 CO o > o o o CO w H CO O 5 H o Q z ^ o cc o d; o K CO O o Q o w o z w W in CQ •< •gSo ^-eT ■2-= S^ I S 5^ ; — 0_>) O Z o E-a. -■= S 0-= S 2 1^ # 5 aj-= ° S S "S " ? ? ^ s 6 as , S.a|J III! ■Is-? . fScca ISsi >?: ! « o c^ „ « o 2 -* ^ = £< =— (T) P?^ m 3 S i *5 > f - 3 5 ~ < r. ■■J ,£ 5.- 3 — < 2 -5 •35 hJ c X ?-. 1. 1— ■ F «u 2 ='t- &r: u >i Jc i a z M X r c (ri E " n 1 >> EC 3 Cj m S = t-ai" tn r^? - i*^ * •=; 5 = - ■=• ' - i-- ^■i 0--t-f i-ift ?■ cm-* o M oS Sit! s-s « vOCICO -t t- C t-f-M •*M CO W ZiJ 2 = II 0 ©00 oooooo 00 0 0 1 « 0 ooo oooeoo 00 0 0 s CL.C 0 000 c 000 00 00 0 to ^1 S3 M f r- » 'f ov •g B-S s s Cl wr- a M *-< MM CJ vO « 2 = ^ 3 » i| i CI COf^CO M^Wa-^O t-i-r eo 9> S OS 10 irjo M M 0 M O) nC f r- CO CO » 3 Ci,c n CO 3 i ST! M — r- ^0 — mtnr* ^M M E-S "t 0 lO M L-: r- z= — M M •a a CO NC>CI r- 0 eo i-i \o 0 rt» C» inej •M .J 0 ^ (XC w B U5M» oecv CTs OS r- a S-fs -M X — r- Ov>OWl-MC?. . c 1 s 3 i-i « B.S ■* t_ i« 33 • CDO I s x=r^ £•3 •t ^ S-s X 0 ir c S' M 1- 3 3 2= M •* 1 •»_ 0 0 ■6 .-. CI ci = V 3 c-l CI CI M t >^CJ £=: •^ c -5T .^ — . ?<" _^ ^_ CI _ ^ 9v s- "3 h C p" Z = M e- > — z u. « "f 0 — = r- •c E^! £= — •< c:: ^ — 'Z'r. ??T? « ^ rt e 't &% ir ON K es US 3 3 2 = « •« 1 'if. II ^2. « s c ^0 c 0 1 i i^' g = c 4 1* 1-s O" ? s s U9 •^ ^ ~" c c c « ! >^= 5 i 1 I'J 1 1 1 1 is i 3 : 1 * J = X S - S ^ T "31 = h.i lis ^^ E i-5 ili^ ^-^lil i^i^iii=5?i= ■3 » i-^ 11* l.l.i?.t i^-B :.^=-" "*S^.3 3 <=£ 5 t- 5 =S O O ro O ^■■■■■■•■■••■•■■•■■•■••■••••a» ^^:^:■^:■:■:^:■:■:■:^:■ ■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:■:, < z Q- ,. O^ O^ CD O) CT) ^ (M OJ CVJ CVJ CVJ Q a> ii — ro (T> zz — m I\vwX\<-7,-. ti',i,.:i:i:i:i:i:i:i:i:i:i:i "I'.''.".'.".'.'- '. '. '. '- '. '. '.) UJ CL xO gen CO uj Q. CO < o o fO o ^ n Q u w O o H O 1 1 r* t- ^ §■? •^ ■* o c^ « X "* » ■* X s- z= s-s c^ rt n a s 2 1 « Cl CI CI S 3s c o © o o o 3 o © 0,= 'i -■ 2 t- f X fn ? o a-S X « rt « 3^ M ■* r_ — z= ? || rt CI •f o Z Tl -»• 'C -* 'J' ^ CI hO 'r a z i a-s r- m in M On o K 2 = CI t "f i i| q If? X i P t 3 CI CI . CI "3 M a.a 1. t- * 1' X B" s-^ t- 1- M X 9' z= = 11 in CO X X I-' ^. ■S s &e: eg so X M « ^ © = 5 11 2 = w* ON Q "■ CI CI T 0 11 _ X ^ 0 X — _^ o © i t 3 CI M CI •« 3 tT: -2 n X ■^ M -*■ O 2 = - ? Q. 2 j= r- o in "* -* s p ei m o V 3 J5 0 2 t-TT ^ 1^ a « X ID 2 = ■5 • 1 .^ ft 3 3 i 2 c 2 ua ;<, C F "S 2 " 2 %. r- a g ?l 3 C/3 0 fi H ? 2 U 1 2 ~" •5'5 £ 3 - 2 -= = L. 7 £ ■ -'■I is ; z.iz P.M -^* N r; •*' < f :i ; = = 1 3£ S-B S-5 2^ =£=^ * = i ^ s = = ■ ^ 7 .? « = « S O QBHSnid 3SnOd9 dO lN30d3d „ri 1- f ? — L- — ^4 CI rt 5 ^•s aWffv ■"* I/: t- c t — ^ ,- rtrt ^ rt o « •B -o ii 6^ 2? L- SC C en rt mm mm ^ e S'^ X XOO"*- vC ^sC^OirtM xr> en « Ov OS ■-fort CI O rt M —1 rt X ^-^ M « \e 1! o ooo ooeooo oo o o 1 § o o o c =: = S 9 S 2 9 2 o© ©© © o o © ."S M nC X = r^ X i: ^M- X — CI in en X « r- -c X C ^ (-: X-* X ff^en « rt -: r^ CI SC « ■g ," 6v- CO ^ ^ r- -C o; CJ OD r^ vd r-' « W --© en o o c ■^ o i-iON sO^OCirt --"O w* in ■■t » » ^1 Cld M »-J *» o C1\C O vcrt x^oo ©>— « o i o 03C oc-^^cs^ CO o © c •u o CJ t^O O- Of^ — ©X ©« ffs o\ ^ ^ \C0O^ rt-eh-c^^« »^ X X ^ M ■^rtTf r-in w*iftt- Tf Tf in in £ •^ 2? 10 O-^nD © X CI — rt o ^o en ffv M 7. 1- 'f'trt -f X -^ ■^ CI CJ CI— w ^cn Cl en ©> S c= 1 6S| M xwMin X t^ <-; N c^ c4 -^ ci -f CI CJ ci ci ff o •3 JS ox M ©in\omxos e s « nC t'- i-i M r- en en ^1 f-ir- in u s U5 •*■*■* Tj-Tf M^CC W n%n ■* ■* o s c V. a 0 C) C — — r- r^ C in ■* ir; O^CI M in en ■^ift-t rrc CI ^ ^ t *t h- r- c ? ^d irtifl-* ■-* « r-; t^ CI r-; cc ift © rt in rt' N en o ITS <= ■* "^ T? ■H cj **■ ci 1- rt *c cv en -* Zi M M -3 S, fS-? s rfO-O -- en m ci o a- ^^cicim" — CI xd in ci in CI o f-— .— .T 32 Ci c ei -?-|-C) s* o- — in© sC X 1^ CI ort o r-r ci o' -* -t r- CIX a^ n zg -t -? i— rt ON o 1 ftS^ « r^ O; o r^ N c' ©' ©' ©' ■-• «' in d in d W «■) c-i — CI W N C^ M ^CI o •P dja win C4 c. 2S c: •iCJ — sO t^ <-< i-i « en en « WCIO ^ -- m Ln rt o^ frt w ^ — ^' — s -■ ai' ^^ 3 CI — cicici- CIM ■^ 6£ o^i-in Off' t- M ZS n en ."2 w ©"♦t^ «« — «»>« «lO N CI 1 tn ««■* oe^'*©rt t- Ot^ N M o C ■g « — CIO- — rt -r Tf ci i/s aeo r- OS « 6 fc ^ff CI 2 2 •« — .-H ff> i «. > — M ^1 -< aauO UUCfaC^US H ! •— -^— 5 1? ■o 1^ ■o 3 8^! ; a c ^ rS t 1 03 < .Uj K s^ "§ "1 c^ (2 h 0 O ? 3^ 2 APPENDIX 841 > n o a o (£ Cd c« O S .= Q o <^ o g O en Z2 I— < -^ Q W o z 5^ b W tc H Z O CC U O o o w o z w z 5S s lT - £ = S 2 lo'-i P ff^ 2 <; 4) u K Si a. o-g M a ^ as c 3-0 q; 0 ^ s = S 5 "2 ' S -^j; *J 3 C3 ^ < 3 , . 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P * ^ ?-| i ■ = ^2^1: ' . - - 5 ^ t ' « 5 i i; E •= " 3f *■ - " = is = >>" J - - > = z :- ?i-^=#:,: ; ? i_ ^ ." — S r c ~ 7 « i - = * i — > '- ^ = = 9 : "5 — X _ 2 » ^ . at— 4 i «- — '-r r Hz I r-'i =•? ■£ E 1 5^ J MSi ? >, T. 3^5 _ '-'_s ; c ~^ i r > i; r ?- ■= * -•^ >» ■- r. _3 U, 2 3 i X - 5 s ? » -■3 « B- £S .5- 2 X 3^ ■■ *"2 : * Z - z> -£= - X 7 *i?— i-^^'-^ i* — - z - ^ U •!• C -fc : APPENDIX 843 I i =,'. i i-^-= 7" 3 >* I K r- ■n H Z a I a -3 W cc Q Z K c« P J U c» IP O Qi a H Q Z U i? H W ffl O Z H K H Z O u w o z w fa z .SuSoZe.-- 3 = ■3= «.=*=.;■ = SI? 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Z Z ** <£ r : 1 o — — — — ^ — Z. 0. z; o *-*■ 1 1 3 ? ? ^ 1 4-» 1 J 1 £ 5^ = j: i ? = 5 £ -s -i -i -5 •n ^ j ^ ^ 5 ■z n 1 ^ t k t. < T > Ui i = = = = > Z" z* : E f 5 3 ;j ? .: 5 1 i^ o ? l5 CJ 1 w w •> I 1 2 5 2 3 3 -3 1 1. 1 ^ C = •— u- h. .z ^ ^ ^ • 1 j ¥ t J _2 N 1 5 s 9 'S' 1 z 1 X •3 i ■1 a 1 1 1 J3 ^a 1 ~ 5 s 'J CI t 1 I 1 ■T. 8 i 1 < 5 1. £ i s < c 3 < e < 2 < s i t 5 jc 1 *- : 1 1 * J c : ■< ■I 1 •3 1 1 1 3 S 1 J m .2 t 5 5 5 .1 1 .1 .1 5 2 i ■H 5 J •J 1 >t -2 1 J J e j c 1 1 J 11 H 1 1 1 -3 •3 ■3 ■3 3 9 k' 5 i ^ 3 3 5* : 3i i : j 1 1 .11 APPENDIX 847 _ b. =»: ^ ^ e =fc iss 'C •* r- = fi © CI »* CX Ti ?i L* r t- t- ^ L*; .Se = ci ti ci ci ti — ■ . f -' \A ^' ^' ^' r* ^ -•' ^ ~^ c _^ i' , J ^ ci J= ggw ■s ^ 11 »fc >«. MM =«» <*! o\ eo ff ?• •* rt ** r- *I ^- NS M © M » Li >» i o T^ » t^ M T? f sc ^ NC o "* CJ © M X ffl © T3 9 2 c d © 2 — ' \» es d d ir t- CI t-*?* d" © e -B u- ■* c» - " r; c^ "^ •' CI — c ci > 0 1 1 S = =»> =». «i« =», — C — ^O s£ C) ■c CI CI r- X 1^ 'S' © c ^ £ « r; I- r: f- C'l 3 t- CI M 1^ «d c^ 1 c U c a-' d ©' © e- r- c c' so ^' cj c' S3 ^ ci ~ ir r^ lO d e- M 8 fe z| w xf. L-^ w w ^e t- ^ Cl 'f « L'S rt t~ 3 X L-i rt « ifl •O a -— ~"~ "= ec f- «• t.- — CC _ _^ ^ X X 1* . ,, „ ^ „ CI _ © 4. if^ o r- sC r- a; CI SC C'l © X r- CI i- CI s •i> III ^i ffi ffi tl t^' M c c e i. ^ ci "f* CI r- ■c ^ .,>' f- ^ ••' ^^-■ ^ a ©■ Q, es o "^ D O F-l u^ C « NO ^ c- i-t « X © X © _^ ^ CI . ^. trt CI IT c r- »- M © r- © © ?- » t- X i 2ok ifl -^' trt ifi -* \A C r y t «' « »- d X d x' d « »' c CI d -^ X d ■^^z — — ~ ! Moisture (percent of fresh weight) in © >c c s — r" tt w S?' «e '^ M i ? 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X - f * - ^ = 1 < rH 1 r e s 2 z z z z i 5 - 1 .2 g .s ^s _ — _a .2 .2 « g P a e 5 *c 5 _t i _C ^ 3 •; 5 .; -g 5 3 ^ = r >■ > > > > _sc : 'i >^ U > > M C > U ^ > 1 E S ^ ^ >■ ^ iS c i >> >, >> >> >. >> >t >) ">. >) ^- £ J X X 1 X 1 X s £ c Z 5 z i z X J; X *? z = il i Z ? c 1 » X J s s T !t OB S t ^ J= ja J J *+ C ^ ja 1 •»-•■ L. u 4-*> b 1 £ £ £ X - c L 5^ 5 -^ ;&. lb ^ -5 j U. -» U. 1 J T 1 1 > * .« u « -5 X_ *£ ^ ^ *£ s_ 7 ai _2 x_ -i 3 £ .£ < =^ = - ^ "^ ? ^ > H H .£ .H > j ? > 3 S > t *3 ^ > > ^ > — .^ -* -* ' >. h. u 5 -7. 7 E : 3 C u u >■ "x s 3 u -r t > c ■1 .1 ^ = ^ .jc ^ ■^ ^ i s " 2 "Z i i :§ ,55 1 „ _ f ^~' ^ .^ i = ■A r = ^ i 1 i > 2 1 J 1 ^ u .= I ,ae 5 1 1 ■j 1 1 f '" 2 5 .1 4 1 1 1 1 i 6 % % II •IJ 5-5 i •* z * > ^ i 1 1 4 7^ 5 1 z 1 a. '1 848 APPENDIX •a V 3 a 'S a o O B o s: o Q U >- 09 Z u H <: o o o tn Q O O "a . ■^^ MM ••. "*. ,^ <«> "^ Hi Si5 f- sO w r* ir: r- X X c ■ m t- « in — ^ — t~ n_ o ^ o -f © * « »n ff> Ll CI -? -- -^ X © ^. r- ■-f en 't SS c d •? ed 1" ci n M t-^ ci ei CI ci Cl ci ci •«" rt : ci d eo "" > "*. "•i ■•■ ^ ^ fc. m rt C-1 © rt M ^ >rt o 0 » in o^ © Cl n « >v 1 2| so -4 u- o- re o «c !> n (7; M o\ © -H r- •^ in in *: 4 ■* X t- ■? o oC h^ sc d * MM ■•i ^ •^ =•= **. 0 ll m C) vO O' o c> f- ff" CI c> 0 Cl © r- 0 ^o ?" • t' in r- ?l s u~ CJ « •? "? C) (- m X CI r- I'j t- >.o r- '. ^. CJ CJ 1 •J X i/i ci ^i c^ « « c^ 3- t-' d X tfi c^ ri — ' w X -^ d ci ^ c '. t^ ci r-' zi irj irt ^ "? -* n n -* •* in "• rt m m m -? in m in ^ ■^ r- en en a, a — ^ "•n o Cl « o © 3> \c n !> © r- r- » en g 1 CI Cl t-; rt q t- CI "^ X «n — — CI X \n t- © C'^ ' ■ X ■o r- 1 t! ■^ is w t) ti M vi ci ci "t p^ © t- ci r-' n ci ci -t in © d en d 2-S-^ -H ^ ci '-« CI •^ M S >* U * u tOI =%. <*i Q p- ^ r- o* C^l t- c^ M © © Sv © r- \C s Cl X '^^■*' ?i CI O o^ t- a -* © ^. &■ -* f Cl •• £ Cl r- ■c ; tn I- X ^ 0 X ro ff^ d •a CS t~ rt _fl c -* in d t- ■0 d d in »n in in c X vC in a- d r 1 U '^ fi w « M s 111 M o ^ X CT^ © X 0 in O^ ^c sa ;-: X — 0 0 '« CJ t^ P5 X « in !> ' n ■* fn in in r- ^ 0* r-* O g_ 8) in CO X t- -^ in in in in m Ln ■* -* 5 . « ifl S3o» •3 c u u ^ ^ ^1 ^ SC U &fi V aj 33 a -= "C -; 2 2 2 s 2 t a a X M r T « ffi 99 a M tc SC .r J 3 -=•3 s 1 *S 1 3 1 -e 1 a. 1 1 = C J 1 J b 1 at 1 i ■* S J 1 ■5 — ■* ll g J- CO E C 6 X CI X 2 X* 2 > 2 X en X X a-; 2 sn X f- i;^ CO en rt M ^ £ ^ X cj f- :: 1 8 a J3 c s <5 t ■j d 3 2 ^ i ^ ^ < -g ■^ J 2 tg ^ £ 0 s S Cb Ex. S 0 o §1 c s> 0 0 c d d ,0 .$ 6 u E u s J" * ■3 s i c 'J 9 o u 6 0 (3 5 <2 * 1 * E e 5 i s s 5 >^ >* >. >. 1 1 i 1 3 3 3 3 H H H r- i 1 i u u u t 5 ? d £ a 6 1 c 1 s « Cfi M !C = 8 c c J 1 1 i .2 •5 0 s a a "E e a a .2 3 J : 3 3 ri S = > .1 be 1 i 1 _> > .1 > > > > > > 5t it U St ^ L i in 1 1 1 = 1 » a z 1 ?? s s ? c = = c = :: = = = = « c 4; c; 4. 1 i .r 5 ^ JS >* V >% .a .s i = s s s i 5 i ^ S ::. e /T " Z. — C s. 1 4~f s ^ J3 i ~ ^ J 1 1 L. i 1 j J » 1 ^ ^ 2 t J t J a j r ; ■Ui • 1 0 > C c ,2 i > t' ; 0 s .— 9 X > ^ '= •< e c •e «> ** e 3 3." ■a c ■3 "a s 5 a Q .0 *3 > S ? •M ? > 1 ii 1 1 a :> 1 ^ 1 i . ? 1 1 < 1 3 i 3 c c > J "i 1 "a '» 1 1 1 - 1 i 1 ; 1 ? i 1 ] ! ~ 1 ■3 1 s 0 ^ 1 ■ i ; -5 1 i I > 1 •1 _5 i ■ s E ■ 1 1 1 E . -a B •2 ae E 2 . 1 ' 1 1 i e .0 5 Qi a .5 ? 1 ; i . ; C i 1 ■ 1 \ i = i N i \ • c j 3 1 i 3 X a t c ! 1 j J e i . 1 E 3 4 1 t J 1 c E •< j 1. i •J Hi 5 - 5 .1 s. 3 c 3 S 1] ; ■ ■■ =• { i < >, 5 3 •i 5 J 2 k 3 5 5 J 4 e ; ; i 1 1 i i. i. h- : ^ ; r^ K * ■^ r ^ r t ■^ •«? •• c -r i* u APPENDIX 849 o U o z w en 13 O EC O Q H b >- P3 z w z o o y (fl Q O O W O O ■ Z O CO O a. o o hj u W K O « -^ ^ ^ ^ -^ 2 E-- K £ X <£ r- 1-4 t- c^ t- in in » t- -* -t C) X "* CI ^ re c» ■^^ e^' ci c-j ci w ci ^ CI .ri cn r- •^ re TJ i^ -* -* — ue ea j= ss- " Ci .2? '5 ^ =* =* =* '^i =*: =*: 1 ii L. CO M 1- « CI OS O* X O re SC >) * ^. © S; rt CO c- *£> es o OS r- OS 'j \h x SO ■B a > U ^^ k. „ir5 '•s ■* =%> =* c a;, SCI t- c c m ur X r- o o CI o k. O X q t— q a CI CI tC Ln -^ o- o c'; sC ■if CI in in in r- r- o^ «' in 1- in o "az M CI ee S-Sj:-? 3 j; M t; r- o o .2 utS.Sf cj c X X p- re X N so »n ci m l!^ c Os' ee' O^' oi "3 S««"*^ t t -i- -# •^■ "^ " in X £ •* in CO in al ^1 i '1 «03 1 U S tot 'i is .1 ? - 1 a C 2 T 2, '5 r J > 2 i 1 1 1 3 1 s ^ "^ h- CI h- c CI r^ ^ s « « ■^ ll § CO eo ee fn 3» c d ^ = ^' t t ~ « S 6 " ci >1 E £ £ _>i u Z O "^ :£ ^ < 5 < < ^ ^ c. ^ ^ >. >. >. >» >i >. t! 0 6 C :S :S Ji = « ^ c c U U c C .§ > c9 > > *3 > 3 "s ' 1 J s C a c c 5 1 — E J .r 1: ^ *» .t: .ti .t: .t; .t: X w CJ ; IE i£ Z Z z Z :« ■s 1 'i s to a s .5 .5 e .? ~ a •z « 'S *c 'E . ^ s "c ^ ^ A « 5 « 5 3 f C .= J a a c « c > > > > > a "; .^ ^ ' > — ^ t ^ Bo'l *>. ">> ">. 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E £ 2 •) = S.2 » « i-n r --Ell rill 111 -■«-♦■* >-++o • 1 850 APPENDIX ig - Q a: r,-, — Q i s: «* ~ o^ Q K C E Ua ;^ o fe O -] S ■^ r^r ^ •^ ■< u g r/) >• c/: OS < .11 X < p - - -J c - 1 .5 -a; .2 if ■n § si S ^^- p4 CI -Hi- M CI ^Cl — - — — s ■< Win d 9- ^N 1 1 d 2 1 1 - -1 o q CI ■=1 8 1 II 1 s ■*sO CI 1 SS "a ■o < .ax S* 1 CI CI \r> \n - CI « *2S- - z:-:: Cl» CI ^ CI CI LI -M? CI o CI f^ — CI CI - E c5 CJ CI CI CI Cl CI fH - — C4 ■c CI - asirtMM CI CI 1 1 1 •c 1 3 8 I % rtoet- g 8 I =11 gd 1 1 «4 ad 1 % III 1 a; s s 3 S" S 1 d d -1 ss - IS II > Ml 1 9 8 - §2 = 1 S i 1 - > < a 1 ] e 3 2 . c 1^ < 1l 3 -3 a 4; In < ^ '3 c u. = 'i i 1. J 1 > 1 i i -^ ill C 1 1 £ e ■< 2 .1 \\\ 1^ "-5 ; ill ill : 1 v. h 1 3 1 e It. "3 i-i "H : l| E 1 u i 11 ij 11 •2 ! e s s a > 1 i 1 s 5 1 3 > 1 •J. 1 c ..1 1 \ 5 S — ^ i1 3 c i '^ "5 ^- 1 5 5 •« 5 1 ■s C £ a c 5 -3 = a: 1 ■c a \ 1 E £ \ e 5 ■< c \ I a £ j ! E a ■S "k a 1 i 1 1 i 2 1 1 • ■a i § 1 APPENDIX 851 ■i 1^ a* 1-2 I s < o - ■^ -fJ - o - c M - a >> M •.f W---H a-* M-« -H N N — (ifl N NCO 3 ^ t>- eo U5 - - - .2 S Jl 3 CO 88 22 1 S d d 8 8- >> 3 a) 1 nil SO u 88 11 8 (9 1 1 II d ^1-1 dd c 9 1 in d d d 1 •* d d 1^ SI si a * 1-2 a -_ rt r- - eo -"S (2 -^ - -HCO N ^ C^Cl^ - ■c — - - a a 3 - CO -^ ^^ t, ^ - - rt^ s '• - - ^-NM be a a - rt -- • C4 n 1 5 V 1 u "S a 3 > 88 1 d 1 1 P2 a i2 II 1 r 1 s 2 888 III 1 IP 0 o ll 1 1 a a 8 88 8 8". 8 1 i II « 8 8 d 888 3 1 1 J 88 IS 1 ^d a , '- 1 li li a a : c 1 i L. i ' C 9J a a c a X c 1 > c s 11 i :3t3 IJ «l z 2l ij a w II 1 s "; J 1 5 5 u t C 1 a c a s 1 -c ■ 2 1 a c ,3 1 1 1 d ] .J .1 1 a > 1 3 "i c bi > : 'c 4. :^ -a Q 1 i 1 1 J 1 s c c C 1 a c 1 i i £ £ ■ b 3 1 1 J c c 1 C £ 1 1 1 1 C n i. 'Z z h c > c > 1 £ c 1 CB > L. s j Z z 1 1 1 1 1 C £ 1 1 a -c u S > a Z J J — 5 'o i i > z 1 £ 1 C 1 c 1 ■j 1 1 c y > £ 1. i 1 e >• li n V ' !l "a J z J a J. 1 3 1 1 '1 a a a. t j 1 i .1 1 > J 1 1 J J c 1 1 . b c E I. a 1 '1 ac a i 1 ■1 'i I. -a 1 3 ■: J f e 1 852 APPENDIX •g U ii ii M ^^vOOO rtn *« s O* CO - e 0^ — •*•»-* ■o UT Mrt t._t- — -.(-C\ - o « — \ew «M eo< d - d d cs cs d 8 a is 1 00 g« 3 g Cl CI tJ 2| d d 1 III do eo §» dad a 3 = 8 s 31 d SS8S8 a tB 1 c i 1. 1 1 1 d a d h. 8 1 1 1 CO d 1 d d od eg » gin g g g geo o «g8S -nNgg ood s a d 88 g a is gg-^ Si 8 88 fit a 1 g 5 25 s 1 1 £ •SI Is a Ii 1 > X 1 1 M 1 1 1 1 1 1 5 1 1 I ■2 "» • 1 1 fi ii 1 4 1 1 1 > J i £ • e 1 1 e j s 1 J 1 1 u s ! 1^ 1 > 1 3 C -5 1 u Z H • s 1 &^ c ■a I N 1 i 1 * J c -Si 1 s c c 1 1 1 1 1 1 1 > c = 1 e X u ■ 2 E 1 1 I 1 1 1 c J 1 J J c; 1 s 1 J i c ■£ .c 1 1 > 2 > s 1 i 1 1 1 1 c § 1 1 1 1 1 i .a * 1 a « u ■i; c a 1 1 % i I 1 1 1 i c s I I z : T C e .J 1 2 3 1 1 ?■ t 3 c ] i .a 1 1 i 1 1 1 C -3 •1 1 i J ^ C II < 1 1 1 CO 1 e ^ > II 1-i ii 31 -1 • : c a •^ 1 1 a -3 3 ra B 3 i APPENDIX 853 a a o O CO Q O O [^ O 05 ■s 2 1^ jl 1-2 3 s irtM « :* Cl N >> t-tM so ^FHM - " eo -- 3 iflM p-l - - •-* CO - Ib II S < O 8 1 do 1 d *3 ■-s II d O O V III 1 \ 1 1 3 3| s s 1 s s d d s 1 ■3 3 11 a n^-* <* Cl-f ^ffv — FH«^ M^O C4 — CO ^ Ift 1 \n ^X ^ ^r-e-i W » 5-J r- c « M rt 0 a i to ^---,.c. i^i-t -* ^rt «lO M^ M s •c a to - « S t- t- MM ^ «=»o«-« 2jg -» — ^o--* S CO in s 5 a s u •c V a •o > S a 5 8 8-1 ir ss IS 88 11 88 SI es doMModd c-i-^dejosrtdddds 8 -1 1 ~ 1 gogo 1 d 2 1 dd gddcid t- ^ 8 1 CO » s a 3 8 8 S 8-1 8 CS eS (9 C90 (8 88 a a an -1 88 ° II 9CS og 1 se d a 1 © |:= d in d =11 d-^^rtd-H dio N 0 d -!• g trt 8 1 •c M s g 1 a T & 1 ?1 2 S = £ 11 g° Ss w ? 2 D c % \ c 1 1 ■ '5 J z J z 1 X 11 ■3_ — ' b a> :| c .^ c §5 c c c \ 1 1 J I 1. > c _t J ' 2 . 3 5 > i b t 1 "i ■IE r .- 2 1 1 > 1 £ i i « ■ 11 1 J £ ■ B 1 z £ k i 3 a ; il 3 > k " 5 E. C ; 1 "£ X c 'Z z s -■|J - i = "3 c: 1 e : c i 1 5 ■1 c 1 1 c h 1 c _£ ^■< 'e = ^-£ ^■= BS n 3 5 1" 1 i C > -C 'C -S --£ S J I 1 ! J 1 I a; u : ; x > J 1 1 J J 1 C e f J ^2 854 APPENDIX m Q O O o o J* 1^ 0 » H 2- K 3 &r 3 < « w C4 M «■* - C->OMW - in - - 0 - 3 - M^4 M - d r^-^ C 3 ift ^ ^M - - " M ^ Mff. iO •II It S < 1 d o SlaiS aooo so g 1 1 88 II S a S V 1 88 II dd 3 © O ffl £ 1 S 8 a 8 1 2| d 3 "a ■El SI II C n - l—W .-H Irt ^^ ^ MCI ■*« « MM 1 M - N Irt^ CI U) — «?4 ^W - M^M M E B 3 - - « ■* N^ N ■** w - 64 M^ - e to M— i W - - -.-, « W - W^N ■c o E 3 ■o > S e d d gd g d 11 i SB IS S8 n d gg II 1 d d ;;j -=8 S -:S».-i o goo go s I s s g III d L. a Si g d d d Si 1 II «d 2 s S I go s a S| 00 t^ 1 II 1 6 d «■! — w § 3 ■ I * i s c J c c i, -" 1 e £ i J E '1 .1 1 '-^ 2 S 3 1 11 e 1 ■ 1 i ■i 1 1 a s ! 7 2 J J 1 I ^ ; ^ >.^ : 1 1 ' 4 r:7 1 s E ' < - < 1 X ■ ■hIIs-: 1^14 5 mil C O 0 «. = ■limine HI ?•- = c Ij S : 3 ( 5 c •IJ \ 31 111 i ill 7.-2 ^i •ifel ir ? Is e'E il ".= ? 3-3 2 S 8-3^ ?- = 3 « s 5 «E il £ ^ £ ■ » c III! 113 APPENDIX 855 1-2 3 -< \nc^ ONON e^ N e^ m h- ^0 - CO - - O t- M M irt » CO - ** -^ CO 0^ - in to ^ - 0^ c 3 ^^CM CO CO - CO wo M - If 3 S ■ofe > a 3 go = _go s d 0) 1 d *5 — 1° a; 1 6d ffld^'d© 8 a s 2 8 2 d 8 g d « 5 1 s i 3 <> — -id do go 1 i eo d 1^ S S 3 "a II a* M PH W « O-H^M-O h- -f — woo CO Of-eo C* CO -f M 1 ej-t Mf^ — r- r:— -^ciM — t- " ?! = co« CO "S 5 s CO S V a a 3 CO — e-ifi--e-jo-N —1 — -t - NP9 - M •* Tl — — ■" r - " ■ ' c-j i « « C a CO « - MCO — ^-N = ?J — «r-ineo»e «eo^ M S" WW 1 1 O •z 1 •o 1 g d d <^■g8 8-; =? © a a a© © do 0^ « gM M «d n S g gn^i g d 2 2 ?^ci a in «o' 1 0 to « ca £ £ a = o so C4 ■* d d CO d deo' ■*iO do N 10 d d B eg o «ooo aoo 1 1 00a ac 1 « g g-^o » W 1 CO d EC C CO d v g — C-l CO -H £ d d d d se cc d d — «^ — « d d d d M d — M dd ^ 1 * i 1 a c ■a a c - IJ "3 c 0 ; §§ g* "c < J c J i 'J i E > ■c X .! c c a X 1 J J ■ e J 1 c ■ S i 2 1 c E .1 3 c < 1 t i t. E " a c '£ >. ll u 03.* X : a.'C -0 <- 1 > 1 1 i a b 1 J 11 y 1 c s c . c 1 L i ■J c 1 > J ' c 3 s c 5 a 'J* c 5 i 1 ! 1 J c ; £ c "t I t < 1 . '} J Hi =211 11 = ^ !*£ = >. : 'si 1-3 J c ^1 J £ c r - c 1 " i > !: 1 L ■a J = t. i 1 i ! . i c i > C c E a "S c c .£ 5 1 ! I'Z c 1 "b c 1 3 J 1 Si's 1^ 1 L i % c E i i ,-c -1 1 ■ ! 1- 1 c J J J 1 J J K. V £ 1 £ £ 3 i 1 c 1 ,1 i i i 1.S 5 'i : : 1 z £ "i = 1 c \ I e in ■J E L. 'J 1 E c 1 1 1. 856 APPENDIX 09 1 1^ II 1-2 1 a es ini-H - t- 1^ CJ - - - — — \C M F^F^-O - a s N-* *♦ r- ■ - ^•^ « M ■11 Jl < 6 g C) CJ cc -* ■ e ui ci M d ; B g -^ d dg s "a g«d M'ddd ; 1 g ==-g 1 °°s ggg CO a CO d u e 3 2S d d : 1 NM dd u g a 3 "a rs < CD ?^ fe.s a» 1-2 a 1^ \D e-; o irt <-i MM 0\ OS i-n-*W sc - ift«eo«0 -ifM N 1 ^^ M - r-l VO F^ --^:ci:-:in3 COM 0) a a a to - 2§ -SSSS'- = N - CO - -Hvcr- 1 MNOClft 0 ^ N r- •* OCNCJ^ -- N 1 f 1 1 3 > c do OMCjci d CO d ci d d d 3; a; o; ej 0 u u u S|SS g«W d 1 -lgg"!° -g 0 a gM-* 0 g g "a 11 o\eo in n -^ r^ CO "*. S ®; *i S .-I pi d i-< d d g d d g 1 d d d g CO Sggg=-i-i -g SSCSSdM OSS a eg CJ gOCO'tf gM gefl'ps^ go 0 d S 8 <=>«, S « S C40 « 0 1 rl Si — ^" d sc d d d d d d ddg«d gg d oe 1 4 X c 1 1 c X a 5 1 1 i c 3 1 1 a. .X 1 1 i > L. ■: .5 u '■ % i. • I > 5 si s •- 1 1 ': e 1 a .J c i 5 J I 1 s 1 i J 1 g-i ^1 i ^2 IS 5| E C ■i t 1 7."l D C 1 i as 1 = c -i a* •5.2 £ J U C3 £ > > J s £ J 3 k 1-= 1 2 c 21 si ii |.h|-S IIS i i-il-feS ij £414 «:|^i i-2 fe :J4|-^S4 sl^i^J S|i.5| s L. < llil APPENDIX 857 u II 1-2 IS 3 m 3 -< -^ m t^ - - - - «Ci - - ■M - tfi '• • •e i--H^M C3\ r- • - : 'm ' • 1/5 M ■ M -a . ;- > ft 1 3 < B dd 8 0 1 n» 8 • ©0 g . 88-i 1; 8 ■0 §« • ;dg6; Id do -tr- od s goo m« g © 8 :d :d ^ 9 d 1 d^ is 1 § o Q O 3 "3 ■< |l a * 1-2 a M^-*W-*CO coin »-» - rt rt ■r \ ;,OM\OWNrt ^ £ i-< 1—1 M 'S - M-^-H CI ■ -? OS « »/: r- CO a a «-* p^fT* ^ - M W M ■ ^ CI \fn-~^x-~ n a M ^ M <^«o ■»*f-F- ^ ■* ■ ^e^irt --co W 1 a T 1 a o « go «o do s 2 8 1 J S ;l ; ■M g-- gci lo 000 gd 1 8 '6 u 1 o go 8'nS S SO a a is 5 i 1 :888 . C9 S CS g,N go s 3 ( t : 3 ; c ! 11 i Si 1 ■ 9 :e . e a Si ■"i ~i ' e : i 5 r •I 3.: 5 1 J J ? ^ 1 ■ - i i ' i i : L ■1 > r -ll" = 13^ • 1 J . c 3 f 5 -■ i. 3^ 1 1 a. : : 0 1 h S S H y = = t s J i -■J * ""J • \ 1- ; s ■ 1 3_ 1 — ' e 4 Sl1 : fti ■ if. ill illllii l|ll||diJ.i||ii o < > X '0 > S (S U 5.. 3 ?ij-2 s-p SSI silsi^rgs 1-11 < 3 U 858 APPENDIX 09 fe.Si 9 ■< . . . .►-(i-tMr^ . .r- - ii* ; i :- : - - • :— - in ^ 3 ^ :;:•-;•; : is : ■ ws * -I M * :— i CI •— !? 1 • •••«-•■ :^ ; li^i ! i - : •O - S If II 1 tic s •< : : § 8 u« • -oo ■ ! Id '. is i iSi Hi |3; :;i= S 1 s = >> ::::§:: . . . . «B . . : : : :-^ : : ■ -co ■ :d '. i|i g 2 iss s S i|5 d T5 3 HniH : :^ : is i i*i ; 1 i i* 8 « s ° 0) = e •s J en Q O 3 < o „ 2- s a ^tn '■ •« CO » -Vh . M ■* -r- • i - - : :- n n • o • o ■ ': " - U 1 >„;_ ■ i—M -r- ■'O ■--'*'- ^ M « ■ h- ■ ■ ^^ '■•^ — i i 2 " S O fc. o i E eg - : : : : ■f-iCOr-i* ^ '« - -- ; ^M<-t rtw-* -^ •^ - ; ■ ■« ■ rt ^ « in o en be a •c a « ■ us -i-tCs ■ «F-i -m ■ ■ -HMCv<-< - : : in ■ • i- - • w ■CO ■ in 1 i 1 u i B a 0 c — iM ■ ■ . o ^' d ! d 1 g :i -. i i! O . .J ! * IS -. IS = :d is i :- ; i S 8 « £ 2 ® =3 i :-.g ;*. is 1 ■'"b '■° ■% : :od g ill! m ■ • d : ; i i -g i- o a .o II i CO g r- ■ o 2 © B S 3 8 : : : ': i •. ; ; ; • ■ ■ ■ g^n>o is i* i iS i° i 1 i 88S SSI \ III :l • ■ M : : d s S i is . . to : :^ ■ •<»« g ■* .1 u a. to .o . a 5 ciM r- ■ ■ s :«d id ' s ; : : ; is^ss 1; :d : is « :S = '. Id ! lA d • 1 i 1 .r 1- 1 1' '. 'a '. '. '^ \ INI ; ■»::-- : -c : ; = ; ; j- n;N!;f ilii|ll.t :*i is iiii 5-!: 2:^ = : K B r!^ -P-J 3 i -1 : s ■ '-— 111 -^ s ■2 Si "is 111 III >, > 1 ■^ f:fr i.ir : 5.- ai - » c . N 1 ii f • ■ X . . ac '5 '. '.'>, J Ill's: M3 i-j 'fir is^" = -■= = ' T is - = E |t 1"- 1 • ; m ill 5 '. >*^ e a c . J e X 2 2. Isi |2 ii :: 5.: 1 ■ ■ J ii = ' u ^ .-55 APPENDIX 859 3 _n '^ c o O , 3 CT« r-M - -- w S ^ ■* - N r- - M if II 3 go 0 1 0 WW dd in 1 8 SO 0 g 0 1 d S-^ 8 i° 1 d 3 L. 1 1 s 1 42 ■3 < Sis II 1-2 C w M coco « =- •^ cjec -M«— _ 1 ^ fC^^lrt - ^^^OMO-<«W W-H-4 — w u a s a en - cr — - CI WOv - -g - -. c x a m (-* M NU3 - eo t- WW - «3 5 c u ■c V B 3 3 a 1 d MM dd s -8 8 a ^w d-^ 8 8 8«. 1 ffl do BO M d 00 gM-" 2 2 £® a s<-ioo ao a WW % -4'd a go B g i: d a ca d © 8 ft EC a tn 1 d 1 d d d^ s r a 1 = s II il ^^ 0. ■c « E 1 1 c 1 J li u li ffl " is |1 U c a i Si C3 J a 1 > n a 7 c a E E ,: 1 \ I 1 e « > 0 1 0 3 X c £ 1 j: ■S s 0 1- 1 J c 3 1 c 1 c E 5 c i £ t IE c £ 1 1 1 C 1 "i 8 "I 1 S E > L. £ 1 X > c. S 1 % ^1 1-1 il £ J i 0 •J E { a £ 1 c 1 3; 1 s s E J 1 e .s 1 £ \ 1 c i c *: t c c 1 1 "a c > .c "5 .; I.. c 1 'i 1 e G c c E E > 1 1 > 1 ; C 0- 3 C £ £ 1 i -D C i 1 E a. •3 C U 1 C •3 Q 0 1 ■"1 i i J s j 'a? m §1 •5-2 3 : 1 a £ c 860 APPENDIX e o U Q O o u. b O 11 1-2 « 9 CJ O NO >> « " ^ M •-fl-l « «^ w^ in ?! - - « a >— C»^ c»^r-- > i i •< iS i > 1 c s , c .1 •~i I E a 0 . = •Id y c 1 I V •9 'H, i ; ■6 j •< f 1 1 1 3 i c < 5 S < c i c 1 .a c 1 ■«; 1 1 j .a " a ■< j 1 * 5 - ft e 1 - < 1 = 11 = - ^ - - - Hi o c ' c 5 , = 1 u' < 1 ■ ^ M i - x^ • .5 1 > -c c J - 1 c « 1 "1 *i 'i * c i < ' C 1 ; 1 1 ] J 1 i .1 i k 4 APPENDIX 861 1 u ■OTa ^ a II 1-2 a 3 < ■M fH -* ift^ t^tf cs "a e-ii-t l-l ■-< «— e-j c-i - - w* 0* - - - N f^M « ^ « — - CNi-" 0^— ^ irt - if Is 1 -< d d d 0£ «8 1 -. s °2 1 -8-1 S ogo g 1 d •>->d 2| 8 a 1 a •a (« Q O £ o 1 dd = s d d a d dd »| 1 1 s s •< 1^ a* 1-2 a « 1 - N N-NM N^ 0^rt «-* 1 fH »-»^« 00 - - **-«»«« -< t- M - C4 6c a - M l-H IS d 1 ■s > 8 1 1 V 1 8 -SS 5 =£2 gs 5S "1 II s en «>> 8 d ".8-8 888 o So a 03 e CO 1 1 Ml a 'C a 1 1 m s s < a o o is c S i -n aj 0 11^ Q 3 T3 c 1 5 (9 a o "c Q ■s c 1 s -c c p "S a 51 eo:r "OX 1 = o a, u CD 1 C _0 c "i 1- , a 1 .2 > 1 s V a 111 «■§§ 111 o 1 c 0. CE X j 1 5 'i I. c. c c 0. li a. V — ~ o a J* u •c u If a G a: ^1 c ca C 1 1 D. 11 il < 1 !l - * e 1 C 1 1 i h Is 2 1 c c 5 s 1 c "F !^ E 1 J 1 3 > C c 1 a c i c t > c 1 >> E ■= = il < < 1 u £ C c c 1 ': J '5 c 1 — , a an c t£ a c — ■*? i i .J II u c ec X ■£ > L. E a £ z 1 C J a t " a. ?E Hx a. c r _5 11 « — a ; I 1 s is £ < 862 APPENDIX a o O c« Q O O b O H CO Cd H CO CO 1 1 3 < o ■^ - M - ~'Z ' n^ in CJ ^ ^^ 04 ^p4 3 — — — t^ci CO t- a^ o^ e-i ma - -* ei — lo -H f i M ^ PI C4 - Is 3 a 8 CO u 3 V 11= i° li 2£ 1 1 III III B S d d o - OS -. g 8 S 8 fj 8 "8 dg d. S 1 S "a ■< ti ^•2 1 eo - 1 l-« iM iH p4 ei 1 =2 - s Ot N - - a - M M -- - 1 a 3 .s 1 1 1 ss s 2 5 1 1 B 3 8 S d S 2 s i 1 a 1 b S is 03 iJ 8 1 8 < 1 c: s 2 CO ( ll S ! c i t c i 1 V u < c a, 1 c c 1 'j 1 ( 1 c c ( if 1 a; ^ .s|j 1 ' ; i 1 S E i 1 c t I V E s e c t 1 1. = 1 k 1 c < I i 1 1 a 3 , '5! hi j 1 1 t 1= ■I- c .11 i < 's > c ■c i 1 c * > i •l > 1 •1 ,1 11 •3s c s ■5 c ;- r- 1 • c 1 i B Z £ S = li 'ft S 1 - St ■9 * ^ < .'> ' I* C a tE 1 e C 'i 'I 1 i i. E S — ' « I T X a 1 * III a, « ffl « q "1 a 8 e ■I C 1 i c . APPENDIX 863 It 1-2 3 < irt ■<* >> "a - « ^MM w - in^ N ^M - i-« - a; a 3 lOi— r- Clin ^-.- Tl—i t- O — JDinCO C^l M - 9 :3 8 a 5 1 Sil S 1 II 1 ii s 1 1 9' •s C 6 o o 3 d go II III gsoo sago o 1 8 s 1 •< 1^ 1-9 a 1 N C4 C4 - - s e 1 1 8 1 1 i 1 s 1 1 1 1 III Hill 1 tic 1 8 a 1 s B 11 A X 3 k 0 T C( Q £ a a c "a iJ Oi 13 C 3 V ■< s &^ O C8 ill go E I a 1 Is is a 33 :° a 0 V 1 a L. c 1 Q C 0 b 10 £ E II 111 Sf.s So: u J ■a > o 1 o ■S ; c I c o .£ a 1 i a ■n I 2 z. 1 c 1 3 a -=£ c i 'S B C : 1 :5 i ; Q •11 3 C 1 1 4 i 5 _^ -3 1 c M i £ 5 1 J ■ s C £ J < "5 -< 1 3 — 1 i : C •fu y ll = ";" - S — \, u 1 1 J 1 1 5 2 ■r i 1 5 4 i * i 5 5 3 a > 1 2 2 .a 1 3 c "= 0, 864 APPENDIX 1 U 1^ It 3 3 •< ■ i-t -r- - MM : : M - :- : :- ; — : w -^xn '.^ ■^ '.tn 1 In 1 1 1 1m 3 — 1 -K ^ ■ • :_ _ „ nr:— ■ ■ ^ CI — . ft L-: — -H 1 Ice^ •II 3 GO 3 :* :* : 1 ::;; Ml >> S : 3 :|| \ :i 111 § 88 :S 1 II :| 2 :S : :S : iHi -n -■- — M = 1 b : :S 8 ".%-.% 8 .a a © SO a a ;i3 is is i; i t D s; 1 ■J !j y . a a a . a a o S 3 -< 1-2 1 a : :" 1 M ■ • • 'a, . -Mr, - ;- r-« '• • ■« ', \ \ O E B 3 to '-2S : ■1^ 1 cecj ■ ■ — -ici -co l^^vO Ed ■ ' IM ^O -• ; - t^^ 1 ; "m ! iw 1 > e ^Vi : 8 : : : . . OS . . . . : * : ; ; 1 : :88 : ..am : :-- ' flj Nl riiin : :^ : : : 1 o o o . 2se° : ;^ : : A J : ".8 : ■ 888 : :- III ::» :l=l B3 < ■a Hill : 8 Mill : :S ; :8 < 3 ; 1?: ell J -^ t|i: a ^2 ■ ~? ; X " = i in ■ o Tit ■ • a ' • e • ■.' : ''^ 5 ■- 5 : - ;'j ; 2 ■ k ii. i|. ii. M - c. - a-* 5 -3 ■ ■ 2'C ■ ■ Mil z c c 2 =-= S^ . — . •. k 0^- 1- Jill: 1 1 '3 1 ! : : i : : ■ ■ Sf - • ^s lis : =i t = § J 3 ill Mi irmili 33151 ?-l i i ia . . . *. • '.5 c ?! "op llll 8s?3 c s 5 APPENDIX 865 1 1? S.2 a* 1-2 1 w rH « >> 3 C^ - - -, ■e e^w ^ - ^^ S3 - N c 3 •-> CI -HrtC)^ ?l c-'i m - - ^ — M || a o 3 3 -< 1 O 1 >> g 1 i II 1 III 1 d g 1 1 i ■a 3 _g a o O Q O o 9 r II 1 O OO C9 d d d 8 O ^3 8 8 3 a 8 8 8 88 C5 a C3 S 3 i i :3 :sij 888 s s s 8 S3 5 to 1-2 a 1 - M et "I a a 3 vO CI ^^~^. — ,, - •*-< a en - C4 f4 fH S 1 1 ■c s 3 1 5 c 1 a S 1 8 M 1- f-H 0) a a s to « 1 gggg ggg III! 2S2 1 II g EC c ■c 1 S 2 i ■5 « 3 3 '■P c a "^ jl 0, s T •l a X « £ I 1 ■c 1 ■fcs a-; 'J ~ i ^2 'II "3. li I 1 J: CO «. ■31: '|g c > 1 •ft 1 « 1 C i z ■ .1 3 < •1 1 J 1 •3 S s I': .5 1 'I L c 4 1 1 be c > 5 ^ s ^ 5 a - -3 t 1 "a 1! > < L. a 1 'i Z 1 ■3 1 P . > • 11 z .11 '5 1 09 JZ C s ■!§ - u — J si V ■ i i i "1 c < 1 "s 1 s a i I 5 866 APPENDIX •i U 1' 1-2 3 - -- ~ P" "a « ^^^ in . - M »-M < 1 1 * k < ea c < 'i J ■; u .2 1 J ■a S ii It 1 w c 3 3 1 • 1 c ■r 1 ! 1 1 1 > CO J 1 J § 1 e 1 1 1 z 3 i •2 J "a 1 1 a 1 1 = = "i 1 1 1 0 J 1! a c Ii § £ U a 1 1 i 1 , >- a a ■z "u > £ s u 'c 3 1 1 "1 1 ?l 11 ll U .5 1 g i *• c5 j e 3 I- c c ■3 i 1 (5 o c 5 1 at •) >^ > 'i e 1 5 J c a i c 1 5 J ■5 3 j 1 APPENDIX 867 □ o u en Q O O tb O H W n u 15 n - - ■'S t--* - M — — -- •M -H ir3«-H M — — \0 M - — M 3 lrtC-1 ■* - ■ -* C) Ov — — PT M fCin cc-i — — — CI rel- et — — — — CI •II if n s < 1 IS 2| S >, a CO 11 II :l 1 III nil 1 s S8S a 1 SS 1 S gg SS i u E 3 S2 1 III 88 : sj u e V u u U u §SSI g o 8 s 1 gg SS :| • 3 ■c a CO 1 1 *• i 11 1= it a i i £ 1 5 5 1 £ 1 a a: 9! i 1 1 >■ 3 L. 5 g 1 -S ; J 3 3 c » 1 1 1 i u 1 > 3 ■o 1 1 i -2 ■ 2' 1" (. z il 1 i 1 t z 1 jl .a s 1 1 1 s 1 5 5 3 C 1 5 _ .1 C j .* J ? s 1 1 1 1 c I .1 z J • E 3 i 1 ■ L ■ a it a. 1 .J; 1 868 APPENDIX 1 U 3 •< U5^ -r- so '. '. '. ■irt9>C4* : ::::;;:; ;- : ; i : ; : : : :- ! a ;- ■ c-1 iciF^ 'm ■ '■•- : ::;":::::: 5 S s 2 a 1 i 3 O > 1 s ■ o - 2 : c ;H;i; : : ::*::::::: W s s 3 01 s : II : 8 : : : a . , . ; - : : : : : : :-88S : i'S^i HmNHil Be i . a :S . a :|S :2 : :; s ; : : :^ : : : : : : Z :l 111 l}l III as I ■is. ■:=■> ■It ^ >»3 i; - 55- 5 = i III ifl S.-o « .c c >. ;2 ;'> : * •t'a - = :; 2 ! « 1 a ; 2 ■ :3 : . ; > ; : ill 2- c 5 .= „= I 1 I ! 8 > I i : : ^l-fi- liiji^- >, 5 = i 0 *. »«- M i M if ; i ; i ; n iiNlMiM = 5 X : ^ 5= : =2 ; .= J:-ir i^-^- 1 a c c 3 u. a N 3 i !■ APPENDIX 869 « li as a - rt P^ - "a in C-l n M w c-i e-j - M - COf^ a s ■-s «g«_. M — (tn ^ M-^-N - ■^ « ^PH -H -. --rtFH if II S 3 < 1 1 8 a is •a d s ca a g s s ill d 1 s £ «8 "I T? a 2-p O = o 2 S 1 d CI — — — ^ d c s d d 0 a a 1 § o en Q 3 ■< t. a »-« - 1 N ift - - - O eg r- o - ff4 -- - -^CJ — — CI CI ^ift 2 cn ■c a 03 - w 3 c S 1 u *c s 3 •o > 5 a 1 c a a s 2 § 1 1 - 1— < S a s CO 1: 1 8 S n g 1 1 1 so g g ad ss eg cm B 1 1 j M 1 ; W w O 3 • ( "i ' i 'c J * i k a > : ■ 1 J = ■; c t. j I 1 s 1.-: ■■ ( 1 c i i -^T" s i •i ■t: :s is z h I 1 s 3^ i 1 2 i "H. ' A] a.' r j * : '1 ■ ■ : i 2" : u ■ii ! » i 1 i ' ; 'i ■ s 0 s i . :;a S 2 S J 1 j 'z a" 1 C 3 u : 5 s "2 i < e ■ £ ;■ i c u < 1 : L j ] 870 APPENDIX « 1^ II s* bC a so - - vC - - — - — « o c 3 -J — K -- —f. - _^ „ -_„ , _- _,, — ,._ If ^1 IS 3 d 1 ■-» o 1 2 S £ III! S8 £2 u a T? 3 CO 3CO 3 3 33 3 II 1 ll! Ilr ^1 °'^ II ="l 9i 3 C O U i o 5 "3 •a 1.1 1 i I -e - - - o a - — M - 1 >- •c Q. to - M be 2 e i 1 o E 3 1 1 s 1 d 8 2 1 eo l-H E d ■s ■ §8 ; ll il n •c c. 1 1 « .1 1 1 l| III . < : \ •1 1 -— i-. ; t ; e e .1 i'l ■ i 5.'' ' j ? ■1 ■ KC -3 s '.^. ; . t '. 3 ■ -'5 i ! § i 1; ^0 ■ c ■J ■■■ I s s _ '3" ; u 0 : : 33 '. e .J -i ■ ; ■' 1 I ' — . 3 ^, ■ 1 ; 1^ ■J: :S ■ 3 5 ; ll i = .2 '■"i-j ;1 ■ ■ e a .!! ;J ■J : S- r -1 * £ 3 ' : J : : 1 J"" < < 1 1 1 ! 1 3 ■ : ■ • s : i L i. J APPENDIX 871 c o O c« Q O O O H t/3 .2 fe.2 < ;- OS - c >• ^^ - - r^ - CI c 3 MM M -" ^M— 1 .2£ 11 S Etc u a 1 d 1 22 1 1 s 1 1 go 8 8 '! s °t 1 ^i 8-8 truce truce truce truce 0.1 truce truce 0.2 truce truce truce 0.3 truce truce O.I truce truce truce 0.6 6.6 truce txuco 0.1 1.3 O.l truce "3 •a < tA ■rs-0 ^ a fe.2 2 c 5 - 1 - w 1' CI 1. B S 3 to ^ w •-4 0O sc — bo a •c a •- - CI Eifi 1 s s > c 5 1 a £ 1 1 1 8 1 g s 8 1 ac 88 a 1 8 m 6 8 S a m •S.2 3 > .£ E S^ 0 Q iE a 5 ca 'a 1 E i -o C P 1 ^ i, ■ 3 • ;^ - 11 1 £ ;5 .2 ■ a ; b-i' J; il H u 1 = 3 '?. E 0 < a; < 3 Q. c K 1 a a .*s '5 < 1 c -c 'no' ;u >. 6 0 a.S « S Is a m •5 E •1 E "5 t O '5 JZ a 5 5 "S c £ at < t < c 2 I 1 1 'Ej 1 a 1 i r 3 0** 1 J 5c .3 ^ z. e c •;s a E « Z 1 i ■3 1 : i e i, 5 'a B iiJi 11 u S X a e i 1 u II e e '1 Q is s "2 5 a c 872 APPENDIX 3 C c o O Q O O O H 00 •3 o il feu js'S S * 1-2 3 ■< rt = - - N 0\ »H ^ « - - - 3 - i-H C4 CO incs « «r- OMrH ifl 1^ -^ ti M rt ^rt M «-< II 3 < 1 1 1 ». 2 ® 8 I 1 ao a 3 8 1 1 8 1 a: e 3 d ca 8 S-i S b— r8«. so a ao^sc 8- 888 BC! a c3 a d 8- 8 I 88 s "3 •< 1^ B 1-5 1 -^ M 1 - - m ^ M s C-l -^ M M a £ 3 »rt ^\o M "K - - M tn P5« 1 to - - fH -* - ^ 1 3 a 1 |S 1 1 1 1 1 1 j 1 8 8 a Q 8 a 8 el a a 8 S 8 I |2 1 1 C 1 1 II c 1 2 1 II B 8 1 B 1 •5 c % C « i 1 c e 0 S 3 1 ^ ill c 2 c i : 1 e ] 1 \ 3 < i C -c .; C C a 5 ii I 1 ] c C a c 1 c 1 * c a ' -1 e i C s 'c C E i 'i •E "\ { c £ * I{ ■|| ■ — -: r 1 u 0. i > j J U a e j 1 c i J •z 1 1 1 "a J .1 c X c J 's s 1 3^ i ' E 1 'C c E 1 C 3 & c 'c ■J c ■c { J 1 ! ■J u 3 u JS u « 8 'S I 1 ^1 ■3 be * i - j ': C e > > 1 5 a a ^ >» II IS r APPENDIX 873 M u 2 •S-o II < - 'ss mm r ■■? - C4 - ;- ^5 'vo s " «^ ^ « ^ CJ 52- - 3 M ^ >« CO C * eOrt ^ffseo ^e»-« MM --mCJ Mi-HCOMfO - 3 s do ^1 a gcddd '■ 8 o I 8 s 888 "1 III >> "a 1 1 ©d 1 g gM g-»-»t-n g adadddci CI >-> d d III! 1 1 w g g =11 O •a o=g«oo 2| g^rt add dd-^ 3d 2|i u u o o o ^ G % ■ -. y |2 see =s="s r 3 g o 0 8 o 5 •OT3 1-2 1 a ^ M o» - 1 M»e ^« w »«» c» - « u V s s 3 to - - ^^ 33" H a •E a to « ^•^^ - «me4 a u 'S i 3 > 5 a 1 1- 1 1 1 II 1 III 1 II III a s s s sgsss a a a a a 8 g g gNNin gg II -•88 = 11 I Co a 1 1 888 s 888 III 1 c a .•1 1 is n a c 1 1 c 1 1 S £ 1 c c 5 -a 1 C 1 i 1 > > 1 £ is E 3 1 a s a J s 2 J. a a s •e 1 1 -c a 'c ii 0. T 3 4 ■it "=! ■of- a 3 cs ■a u ■£. i'5 3 a 1 .S E -a j c i a 1 j e a ■j £ a 1 E 3 a 1^ 1 o a 2 = > > 5 a "3 1 2 "S. si ll C o a:^ Is c go 1 C3 tf i 5 ■3 a; ^ a ; 5 c 1 'E <; 5 J |i ■<: 1 E is S 3 i c c s. E > a 1 ■3 a 1 3 2^ 1 e a -< - " O a:2 J 1 i 1 874 APPENDIX c O Q O o b. O en O c« >■ en pa < l1 ii 3 3 ;- >> "5 - -^ — - - ; 10.-1 ! « F^ * :--::: : : :- : " 3 ;- : : - :- - - ; -^ M • ■- - ; 3 ■ ■ :- : 0 ■II 11 1 GCi 3 < "3 § SSSS S 8 a m a a a a a do .go 1 ;; S c 3 id 2 £ 'JO a ii : S : a . * : 1 1; i II n :; 1 : HI 1 : ".g :S : =1 :S d : it 1 a c •c 0. CO i ;i g 1 : a; 0 u s5 s i • E S' 1' s s ■E i = 2 ■? c 5 '■ ! ; ?a - ill "^ «^ U Ms m 1 : . i, t : 3 : - 0 ; III = 'C J' ■ *■ J ; -a III -. t. *< tli - lis 1 li^ t i.F-|t i|r- Ml 1 f=t -i.l it lis 3 c^ ■ r 1 1 .0 :; 2 \2 - 3 I"? . c ■ i" 1 ii APPENDIX 875 TABLE 174. SEASONAL DISTRIBUTION OF PREDATOR STOMACH SPECIMENS EXAMINED Great horned owl. . . . Barred owl Snowy owl Goshawk Cooper's hawk Sharp-shinned hawk. Red-taileii hawk Red-sliouhlered hawk Marsh hawk Red fox Gray fox RonaparU? weasel. . . . New York weasel. . . . Weasel sp Mink Skunk Raccoon House cat Jan. 14 3 .3 1 1 1 3 2 3 12 3 2 Feb. 1 4 13 1 12 1 2 1 1 2 14 1 1 April 16 .5 3 2 4 10 23 17 4 May 21 6 2 3 8 19 13 7 5 1 3 12 2 June July Aug. 2 1 2 15 4 4 Sept. Oct. 14 1 13 6 3 28 1 i 6 '3 Nov. 31 3 1 11 1 Dec. 11 1 6 4 4 3 10 1 4 3 Total lis 20 6 14 41 S3 76 45 3S 134 20 25 12 20 12 88 39 18 TABLE 175. NUMBERS OF \ AlUOUS PREDATORS TAKEN Dl RING CONTROL EXPERIMENTS ON CONNECTICUT HILL ARE.\— iy30-1935 Number taken Species 1930-19.11 1931 -1032 1<)33-1M4 1934-1935 Complete control sub-area Selective control sub-area Mammals: Red fox 8 2 2 31 U '.i 25 25 M 1 5 U 1 2 *> '3 5 24 ■\ I 11 13 60 9 6 1 35 12 •". 4U "T 2 i 6 8 7 1 30 31 10 4 lot i 19 6 1 4 16 23 37 1 1 1 11 .3 4 13 10 1 1 1 2 lit 8 3 12 16 16 2 5 .3 3 '7 111 (iray fox Iloiiapurtc weasel New York weasel Mink ^ Skunk 4 2t Woodchuck Birds: (Irt'jit horned owl liurred owl Screech owl Cooper's luiwk Sharp-shinned hawk Red-tailed hawk ;. . Red-shoulderod hnwk Rou^li-lef^gcd hawk IJroad-winged hawk Marsh hawk *Not trapped in 1931-1932. tThe two species of weasel were not distinguished after 1931-1932. Specimens are listed under the pre- dominating form. 876 APPEXDIX TABLK 176. Qt ALITY OP i:\ IDENCI-; \SCUIBi:U TO DIAGNOSES OF BROKEN-UP GROUSE NESTS FOUND BY THE INVESTIGATION AND ATTRIBUTED TO PREDATION 1929-1942 Species Foj Weasel Fox or weasel Skunk Raccoon Dog Woodchuck Red squirrel. Reil squirrel or chipmunk Chipmunk or mice Opoeaum Bobcat House cat Crow flrent horned owl Cooper's hawk Hawk sji Hawk or owl Pheasant Black snake Sub-total Predator undeterminable. Total Quality of evidence E^xcellent 40 17 22 5 10 6 1 ii 4 1 Good 74 17 i9 16 11 Fair 56 15 40 20 7 5 Total 170 49 40 61 43 19 9 10 3 2 1 1 2 21 8 2 1 IS 4 463 88 551* *The total number of records used in this table differs from that indicated in table 35 because riest.s not comparable for survival calculations could never- theless be included here TABLE 177. QUALITY OF EVIDENCE ASCRIBED TO DIAGNOSES OF DEAD GROUSE CHICKS FOUND BY THE INVESTIGATION AND ATTRIBUTED TO PREDATION —1930-1942 Species Quality of evidence . Total Excellent Good Fair 2 1 7 6 1 3 9 7 3 5 9 io 1 *i 1 21 Hawk or Owl Crow Fox 16 Weasel Total.. 20 19 25 64 TABLE 178. QUALITY OF EVIDENCE ASCRIBED TO DIAGNOSES OF DEAD ADULT GROUSE FOUND BY THE INVESTIGATION AND ATIRIBUTED TO PREDATION —1930-1942 Species Quality of evidonco Total Excellent Good Fair 4 8 2 ii 35 11 3 i o| 'b 1 1 26 98 25 5 1 55 4 2 i 25 127 90 10 6 mi Goshawk . 111 Great linrnril nwl or ^'o^iliawk. . . . Ceriment Year 1931 1932 1933 1934 193S 1936 1937 1938 1939 1940 1941 1942 Incubation Natural: X X X X X X X X X X X X IJen Artificial: X X X X X X X X X X X X X X X X X X X Ilutchability of wild vs. artificially incubated X X X X X Ilatchability of artificial incubators vs. bantam Brooding Natural: X X X X X X X X X X X Hen Artificial: X X X X X X X X X X X X X X X X X X X X X X X Litter X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X "x" X Rcorinp: X X X X X X X "x ' X X X X X X X V X HoIdiiiK o'kI wiiitRring: X X X X X X X X X Shelter X X X X X X X X Breeding and breeders: X X X X "x" X X X X X X X X X X X X X X X X X X X "x ■ X X X X X X X X X X x x x X X X X X X X X X X X I X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X "x ■ X X X X X Egg production from wild vs. hand-reused breeders Feeding: X X X X X X X X X X "x ■ X X X X X X X X X X X X X X X X X X X X X X X X X 878 APPENDIX en CO W O O ^7 Q D <^ ^s OQ r u 5^ >% «g =^z ig E- o y ■< 3 a, Q - O £ O ; O u ; a, CO •J pajB3j jac^nf^ las »|r;ji}j jo M o> irt t- — Irt t- — M c- ro ^ r.c--sCMr:a--^ JO J3quin\j JO jaqmn^ JO juq{un\[ pat|3)ei( pnoi JO pajB3j jaqmnjtj 'jas a|pjaj jo JO juqiun\] JO jaqoinf^ p!Q| s3^ JO jaqmnj^ 6jup33jq JO Jaqnin]«j c ifl cit) — X — ;^ L* c>i U5 w sea — o>c-f — -^r^ot-eo — ci^x>cr-cj — — — — c"* — — — — i-r-iftw©«r^ cj c* ^c c f- fC CI — — ■-* CI CI -^ X t - ■— 3- n f- t- ON r- — "O^c-^x-cci-t-^ot-c*: CI :CsCC*rc ci -H — — JO jaqiun\] paqo^sq \xi\cf\ jo pajcaj ^uaaja^j pajnaj jaquinfyj las apiJaj jo paqa^oq luaajod JO j»qiun[^ las R>f»a JO jaquiiif^ JO jaqainf^ c?^cc^x-cioi/;>flr-t-M sCCliflCiilrtOCCJ-fXX cii-!-:x — ci-ctccciMO — CI X r: a- CI CI •? -t r^ M -c c^» CI c( X <— •— in M « X M(MX w % ££ hX - H fe 2^^ ce s In z. 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CO CO a JiMJZJi V v « sec SCO m u u La Iri b (S s a aoecao 3 3 3 CO CO CO '-H.MCicicicicn«M««Tf-f'^irM,'^ift\e cocorte*J!OcOrtMcocOrt!*jcoJOJ^f^f^f^ \o^er-r-eewxsoCT«avo-Hcici fOTCfOfOf^f^r'«rtrO"(?"*»*-* 1 a o I 1i t5s •Z 0 .■=■0 til== »=o s'H 5 oS— 3 u ^ o E = ^^ S.r i E u " ^ *3 2.S 1-0 a c^ -=: :; w C 3 u -■-,:< a CO s " 3c -03 3 = " 3 3 " o--— , — —._• a; i» .^ — r > r - a' 882 APPE\D1\ lABLK WW. HKCOMD i)V PIU)DrC'H\HV M:\I{ liV W.KW ()\ TIIK COWKCTICri' HILL AND ADIUONDACK. STL DV AREAS— 1930-1912 Year Breeding population Adults present May 31 Potential chicks Chicks hatched Chicks sur- viving to September 1 Adults sur- viving to SfpU'niber I September 1 population Overwint<"r loss Connecticut Hill 1930 92 142 225 283 199 236 170 123 148 177 184 132 167 75 137 208 261 176 245 159 113 142 167 171 11!! 1 |i) 529 848 1334 166R 1069 1 30» 1 009 8U4 912 1004 1178 804 953 200 461 396 330 529 365 33» 474 625 264 431 339 85 1.38 269 257 101 167 158 178 231 HI 173 76 76 138 196 197 16:! Vtn 1 II 115 1.36 163 165 123 127 161 276 465 274 420 30(1 311 273 334 394 276 300 203 19 1931 51 1932 182 1933 1934. 164 1935 1936. 1.30 188 1937 125 1938 157 1939 210 1940 144 1941 133 1942 Adirondack 1932 40 36 24 21 21 22 25 (35) t (28) (29) (27) * 187 176 110 99 1.54 1.32 143 {187)t (143) (154) (143) 83 109 90 60 80 100 70 112 92 70 86 38 25 44 48 41 17 40 39 28 37 32 26 29 16 16 18 25 30 24 25 23 70 51 73 23 64 59 42 70 63 53 60 34 19.33 27 1934 52 1935 o 1936 42 19.37 1938 . . 34 (7)t 1939 (42) 1940 . (34) 1941 (26) 1942 ♦Spring surveys on the Adirondack area were not carried on during the latter part of May and, therefore, did not permit a May 31 pQpulntion estimate. tFigurcs in parentheses ( ) were computed from the ratio of the bropding population to the September 1 population for the years 1932 to 19.18 since spring surveys were n«)l conducted after the lutter date. TABLIC 18.>. SKASONAI, POPII.ATION KSIIMVIKS KOI? TMK VAIUOI S r.oMPXnTMKXT.^ OK INK (:()NM:(y|l(.l T hill STI DV AKKA 1'):{(1-I'iI2 Compartinrtit 1931 1932 i, w u _2 ; S * * & * o o o o c X XX XX i' i" b '' ■'■ t: C "■ "' "• "■ -.-•■>->»>■>• >■ >• >» p.W8 i»:o|0 I a s o = c r: pinioil jod enn:(il 1 o o § if £ i~ — — oci — ri-r-rT )Ud3jad 'uijoij JO ISO) auiiino aScjOAY UUU SjU&ib^! ~ OO 00 00 S< CV pauvaga poos .C^i^uun^ ^pdjinbaj ISuiuvap JO adXj, 03>moC00C2Ci; 1- — 000-roc 3 0 o_ m c C) = o c m c m o^ 3 o^ c: c c o 5 r 7vi ^i o ac cim' cm -r t~ m*m m ■^ tn t^ *n c" ♦+*J '-^ 1; I" "5 ' ■'• m d> 6 S ii =■! ^ — -mo» lom =ccc* = c "^000 -; — ^e^^ — — OO — cic^ — — mr;- "• c**0"i 2c!irim:iio^:imoooofcocsi— mti"*" «««cj 5 5 — 00" 0 0 94 — C4iOtOC^C^C4C>4C4M'VC4dC4C>IC>tC4 joas jaa|]oo I ^ O O -^ O O 01 auiii iC'-j -!; ■! ^ J; .J* . i_ -. ^^ -. - ts^X'tKOxcC xcccooiStoOx-^ xcn<= S-- LITERATURE CITED 885 LITERATURE CITED 1. A.G.C. 1883. The Fluctuating Grouse Supply. Forest and Stream 21 (10) : 185. 2. Abbott, C. C. 1895. The Birds About Us. J. B. Lippincott Co.. Phila. 3. Abbott, Clinton G. 1918. "Biddy". The Conservationist 1 (8) : 115. 4. 1920. Billy and the Steel Mule. The Conservationist 3 (10) : 155-157. 5. Ackert, J. E. 1931. The Morphology and Life History of the Fowl Nematode Ascaridia lineata (Schneiderl. Parasitology 23 (3) : 360-379. 6. Ackert, J. E. and G. E. Cauthen. 1931. Viability of the Eggs of the Fowl Nematode Ascaridia lineata (Schneider) Exposed to Natural Climatic Factors. Jour. Parasitology 18 (2) : 113. 7. Ackert, J. E. and C. A. Herrick. 1928. Effects of the Nematode Ascaridia lineata (Schneider) on Growing Chickens. Jour. Parasitology 15 (1) : 1-13. 8. Adams, W. S. and S. B. Nicholson. 1933. Svmposium on Climatic Cycles. Proc. Nat. Acad. Sciences 19 (3) : 371-375. 9. "Adrian Ondack". 1881. Deer and Partridge. Forest and Stream 17 (18) : 348. 10. AiDRiCH. J. W. and H. Friedmann. 1943. A Revision of the RufTed Grouse. Condor 45 (3) : 83-103. 11. Allee, W. C. 1938. The Social Life <.f Animals. W. W. Norton & Co., New York. 12. Allen, A. A. 1927. The Autobiography of a Mother Grouse. Bird Lore 29 (6l : 444-446. 13. 1928. Diseases of the RufTid Grouse. American Game 17 (2) : 29. 14. 1929. Ten Years' Experiments in Rearing Ruffed Grouse in Captivity. Trans. American Game Conference 16 : 3-21. 15. 1934. Sex Rhythm in the Ruffed Grouse (Bonasa iimheltus) (Linn.) and other Birds. Auk 51 (2) : 180199. 16. 1934. Breeding .Seasnn Behaviour of ihe Ruffed Grouse. Trans. .American Game Conference 20 : 311-322. 17. .\llen, a. a. and \. O. Gross. 1926. Ruffed Grou>e Investigation, 1925-26. .American Game 15 (4) : 81-84. 18. Allen, E. A. 1934. Eiineria augiisla sp. nov. and Eimeria bonasae sp. nov. From Grouse With a Key to the .Species of Eimeriit in Birds. Trans. .'\m. Microscopic Society 53 (1) : 1-5. 19. .\llen, V. C. 1936. Ruffed (;ruu>e Shooting. Game Breeder and Sportsman 40 ( 10) : 216 217. 20. American Ornithologists' Union. 1931. Check List of North .American Birds, Fourth Edition. 21. Anonymous. 18.56. Pheasant, or Partridge. .Moore's Rural New Yorker 7 i48) : :J85. 22. Anonymous. 1875. Trapping Ruffed (Jrouse. Forest and Stream 5 (16) : 243. 23. Anonymous. 1878. (Note re. ruffed grouse in Texas. I Forest and Stream 10 (14) : 256. 24. Anonymous. 1897. .Amending the Game Law of New York. American Fiehl 47 (12) : 267. 25. Anonymous. 1915. Problem of Ruffed (Jrouse Breeding a Difficult One. Bull. American (Janie Protective Assn. 3 (3) : 5-6. 26. Anonymous. 1933. Porcupine Eats Grouse Eggs. Pennsylvania Game News 4 (3) : 5. 27. Anonymous. 1934. Grouse Fights Fire. Pennsylvania Game News 5 (4) : 7. 28. Audubon, John J. 1831. Ornith(dogical Biography. .Adam Black, Edinburgh. 29. 1856. The Birds of America. V. G. Audubon, New York. 30. Bailey, H. II. 1941. An Undescribed Race of Eastern Ruffed Grouse. Bailey Museum and Library of Natural History Bull. 14 : 1. 31. Baird, S. F., T. M. Brewer and Robert Riih.w.w. 1874. .A History of North .American Birds 3. Little, Brown & Co., Boston. 32. Baird, S. F., J. Cassin and G. N. Lawrence. 1858. General Report Lpon The Zoology of The Several Pacific Railroad Routes, Birds — in U. S. War Dept. Report of Explorations and Survevs in 1853-56. Serial No. 799. 33rd Congress 2nd Session, House Exec. Docs. 11 : (9). 33. Bangs, Oi tra.m. 1912. A New Subspecies of the Ruffed Grouse. Auk 29 (3) : 378-379. 34. Barcer, E. H. and L. E. Card. 1938. Diseases and Parasites of Poultry. Lea & Tobiger, Phila. 35. Bartlett, Mott L. 1924. Ruffed Grouse ami Fruit Trees. Am. Game Protective .Assn. Bull. 13 (1) : 19. 886 LITER -ITU RE CITED 36. Bartram, William. 1791. Travels 'riir : 8-11, 32. 59. 1939. Some Characteristics of The Periodic Fluctuations In Abundance of Ruffed (irouse. Trans. Am. Wildlife Conf. 4 : 478-481. 60. Bl/RNiixM. John B. 1918. Why (irouse Are Scarci-. Am. (ianie Prol. .\ssn. Bull. Till : 4-7. 61. BiHTCii. V. 1927. Near Cannibalism In a Buteo. Auk 44 i2l : 248. 62. ■■Byrne." 1882. What l)eslio\. Tlie Kulled (;rouse':' Forest and Stream 19 i H : 66. 63. C.D.P. 1887. Fiehl. Marsh and Scrub. Fore-l ami Stream 29 (241 : 466. 6L Cxi.riiVM. P. A. 1934. Experimental Studies On riie Transmission of Gapeworm i.S v h a' " '" " s irnchni) 1!> Earthworm-. Proc. Roy. .Sic. Series B 11,S ( B791 I : 18 29. 6:.. Cviii'ENTER, F. H. 1887. Bird of Northern Maine in Fall. Ornithologist andtJologist 12 (111:188. 66. Cmii'Entkii, T. M. 1924. Tables, Factors and Formulas for Computing Respiratory Exchange and Biological Transformations of Energy. Carnegie Inst, of Washington, Publ. HO.^a : 121. 67. Cm \i)ii()( k. T. T. 1911). (;roi]-i- Crop ln\estigalion. X^'isconsin Conscrvalion Bull. Tt i3i : 2.321. 68. CiiM'MW. FiiWK \l. I'^()9. ll.indl k ..( r.inU of Ka-li-ni North \in.-rica. D. \|.|.l.i,.n & Co.. New '(ork. 69. (ImiK, L.. .'^. Leon MU) and t.. l!i \il'. I9.U). 1 iglit ami Heproduclion In (ianu- Itinl-. Siieiice 83 (21.50) : 268. LITERATURE CITED 887 70. 1937. Liglil and Tlii- Sexual Cvcle of Game Bir(l~. Science 85 (22051 : 339-340. 71. Ci.ARK, M. W. 1879. Domesticali-il Kiiffe.l (;riiuse. Fnrest and Stream 12 (17) : 326. 72. Clarke, C. H. D. 1935. Blood Parasites of Ruffed Grouse iBonasa umhellus) and Spruce Grouse iCanachites canadensis) With Descriplii>n of Leurorytnznon honasne n. sp. Canadian Jour, of Research 12 (5l : 646-6.50. 73. 1936. Fluctuations In Numbers of Ruffed Grouse Bonasa umbelliis (Linne), With .Special Reference to Ontario. Univ. of Toronto Studies Biol. Series Bull. 41. 74. 1938. Organisms of a Malarial Type in Rufled Grouse With a Description of the Schizogony of Leucocytozoon bonasae. Jour, of Wildlife Management 2 (3) : 146-150. 75. 1940. A Biological Investigation of The Thelon Game Sanctuary. Nat. Museum of (Canada, Dept. of Mines and Resources Bid). 96. 76. Clayton, H. H. 1943. Solar Rilation- To Weather and Life. I. Clayton Weather Service. Canton. (Mass.). 77. Clements, F. E. and V. E. Shelkord. 1939. Bio-Ecology. John Wiley & Sons. New York. 78. CoNOVER, H. B. 1935. .\ New Race of Ruffed Grouse From Vancouver Island. Condor 37 (4) • 204-206. 79. Connecticut. 1924. Biennial Report of the Slate Board of Fisheries and (iarne. 1922-24 : 67-68. 80. 1938. Biennial Report of the State Board of Fisheries and Game, 1936-38 : 18. 81. CoNNELL, F. H. and H. M. Doremls. 1937. Kndopara-itisn\ in Ruffed (Jrouse Near Hanover, New Hampshire. Auk 54 (3) : 321-323. 82. Cooke, W. W. 1888. Report on Bird Mipration in the Mississippi Valley. U.S.D.A.. Div. of Economic Ornithology Bull. 2. 83. CooI'ER, J. G. 1870. The Fauna of Califcirnia and Its Geographical Dl-lriliution. Proc. California Academy of .Science 4 : 61-81. 84. f:oTTAM, Ct.AIlENCE. 1935. Economic Ornithology and the Correlation of I.ahoralory and Field Methods. Wildlife Research and Management Li'affet BS-30. (mimeographed). 85. CouEs, Elliott. 1903. Key to North American Birds 2. 5lh Edition. The Page Co.. B. Iliallh Repiv. ,^6 : 1905-1906. 88. Cram, E. B. 1927. Nematode Parasites of Birds. V. S. Nal. Museum Bull. 140. 89. 1928. Nematodes of Pathological Significance Found in Some Economically Important Birds in North America. U.S.U.A. Tech. Bidl. 49. 90. 1931. Developmental Stages of Some Nematodes of the Spiruroidea Parasites in Poul- try and Game Birds. U.S.D..A. Tech. Bull. 227. 91. 1931. A Comparison of Internal Parasites of Ruffed Grouse of Labrador with Th : 137, 148. 106. Douglass, A. E. 1928. Qimatic Cycles and Tree-Growth. Carnegie Inst, of Washington Publ. 289 (2) : 1-166. 107. Dunsinc, W. J. 1918. The Scarcity of Kuffed Grouse. American Field 89 {6j : 131. 108. Durant, a. J., and E. R. Doll. 1941. Ulcerative Enteritis in yuail. Univ. of Missouri Agr. Exp. Sta. Res. Bui. 325 : 5, 7. 109. DwicHT, J., Jk. 1900. The Moult of North .\merican Telraonidae iQuails, Partridges and Grouse). Auk 17 (1, 2) : 34-51, 143-166. 110. E. S. K. 1884. Massachusetts Game Prospects. Forest and Stream 22 (12j : 227. 111. Eaton, Elon Howard. 1910. Birds of New York. N. Y. S. Museum Memoir 12 (iK 112. 1926. Game Enemies. Game Breeder 28 [4) : 127. 113. Edceworth, F. H. 1935. Cranial Nerves of Vertebrates. Cambridge Univ. Press, England. 114. Edituk. 1874. [Comments Following Article "Grouse and Woodcock."'] Forest and Stream 2 (2) : 22. 115. Editor. 1925. Interesting Grouse News. American Field 104 (41) : 404. 116. Edwards, Gcobce W. 1754. On the Pheasant of Pennsylvania, and the Otis Minor. Philosophical Trans. Roy. Soc. of London 10 (63) : 450-452. 117. 1758. Gleanings of Natural History 5. London. 118. ElFRiG, C. W. G. 1910. Stomach Contents of Some Canadian Birds. Ottawa Naturalist 24 (1) : 18. 119. Elliott, Daniel G. 1864. The Game Birds of the United States. U.S.D.A. Annual Report : 363-364. 120. 1897. The Gallinaceous Game Birds of North America. Suckling & Co., London. 121. Elliott, Robert. 1906. Extracts From the Diary of the Late Robert Elliott. Ollawa Naturalist 20 (6) : 120-126. 122. Elton, Charles. 1924. Periodic Fluctuations in the Numbers of .Animals: Their Causes and Effects. Br. Jour. Exp. Biol. 2 : 11916.3. 123. 1927. Animal Ecology. MacMillen Co., New York. 124. 1942. Voles, Mice and Lemmings. Clarendon Press, Oxford. 125. English, P. F., and L. J. Bennett. 1942. Food Habits of the Gray Fox in Pennsylvania. Penna. Game News 12 (12) : 10-22. 126. 1942. Red Fox Food Habits in Pennsylvania. Penna. Game News 12 (11) : 6-7, 22-23. 127. Ebrington, p. L. 1934, Vulnerability of Bobwhite Populations to Predaticm. Ecology 15 (2) : 110- 127. 128. 1937. What Is the Meaning of Predation? Smithsonian Inst. Annual Report. 1936 : 243-252. 129. 1937. Winter Carrying Capacity of Marginal Ruffed Grouse Envimnmenl in Norlli- central United States. Canadian Field Naturalist 51 (3) : 31-34. 130. 1942. On the Aiialvses of Productivity in Populations of Higher Vertebrates. Jour. Wildlife .Management 6 (2l : 165-181. 131. Errington, p. L., F. Hamersthom and F. N. Hamerstrom, Jr. 1940. The Great Horned Owl and Its Prey in North-central United Slates. Iowa .State College Res. Bull. 277. 132. EWBANK, Ernest L. 1907. The Ruffed Grouse Scarcity. Forest and Stream 69 (24) : 936. 133. Farrow, E. P. 1925. Plant Life on East Anglican Heaths. Cambridge Univ. Press.. England. 134. Fellows. J. Otis. 1880. Weight- of Ruffed Grouse. Forest and Stream 15 i20i : .389. 135. FiNNEGAN. G. P. 18W. Parlridgi- and Woodcock. Forest and Stream 53 (10) : 185. 136. Fisher, A. K. 1907. Hawks and Owls From the Standpoint of the Farmer. U.S.D.A. Iti.d. Survey Circ. 61. 137. Fisher, Lee William. 1939. Studies of the Eastern Ruffed Grouse in Michigan ( lionasa umhelliis umbellus). Michigan State Coll. Agr. Exp. .Sta. Tech. Bull. 166. 138. Fisher, R. A. 1937. The Design of Experiments. Oliver & Boyd, London. 139. 1941. Statistical Methods for Research Workers. Oliver & Bovy. Manly. 1899. Maine Partridge. Forest and Stream 53 (12) : 333. 183. 1905. An Old Cock Partridge in Charge of a Brood. Forest and Stream 65 (2) : 27. 184. Hausmann, F. 1908. Grouse in Chenango County. Forest and Stream 71 (17) : 656. 185. HELLEn, V. G. 1938. The Ch.mical Composition of Oklahoma Rainlall. Okla. \gri, Kxp. Sta. Tech. Bull. 1. 186. Hkilmers, Hlmiv, 1910. A Study of Monthly Variations in the Nutritive \'aluc of Several \^inter Deer Foods. Jour. Wildlife Mgt. 4 (3) : 315-325. 187. Herms, W. B., and C. G. Kadner. 1937. The Louse Fly, Lynchia fusca. Parasite of the Owl. Bubo virginianus parijxcus, a New Vector of Malaria of the California Valley (,)uail. Jour. Parasi tology 23 (3) : 296-297. 188. Hehsey, F. Seymoi «. 1923. A Nestling Red-shouldered Hawk's Hearty Meal. Auk 40 (4) : 693. 189. Hktrick, B. D. 1935. Grouse are "Scrappy" in Butler County. Penna. Game News 6 (4) : 15. 190. Hewitt, C. G. 1921. The Conservation of the Wildlife of (Janaila. Chas. .Srilu»'r"s Sons, New York. 191. HlCGlNSON, F. L. 1630. New England's Plantation. New England .S>c.. New York I 19;«)K 192. Hodge, C. F. 1905. The Drumming Grouse. Country Calendar 1(7): 6406-14. 193. 1906. Nursery Food of the Ruffed Grouse. Country Life In Amirica 9(6): 746. 748. 194. Holland, R. P. 19.37. Disease Or Vermin? Field and Stream 41 (9( : 1."). 195. Hoksfall, M. W. 1938. Ohservations on the Life History of Raillieiilina erhinotiolhrida and of R. tetragona. Jour. Parasitology 24 (5) : 409-421. 196. Host, Per. 1942. Effect of Light On the Mcudts and .^^equencc of Plumage in the Viillow Ptarmigan. Auk 59 (3) : 388-403. 197. Howard, H. E. 1920. Territory In Bird Life. E. P. Dutton & Co.. New York. 198. 1929. An Introduction to the Study of Bird Behavior. Camhridge Univ. Press, Eng- laml. 199. lldHKii. A. B, 1937. Morphogenesis of the Shoulder Architecturi': Aves. Auk 54 (31 : 364-375. 200. 1938. .Muschs of the Avain Hip and Thigh. Auk 55 ll» : 71-88. 201. HiDsoN, G. E. 1937. I^ludies on the Muscles id the Pelvic Appenilage in Binls. AnuTican Miilland Naturalist 18(11 : 1-108. 202. Ill KK, C. G. 1932. Studies on lliii-innproiriis of NhMiiniug Dovi-s. Am. Jour. Ihgienc 16 1 2) : 618- 623. 203. Hi NTINCTON, Ellsworth. 1931. The Mateinak (!onference on Biidogical (!vcles. ."science, n.s. 74 (19141 : 229-23.5. 204. Isini, N. 1942. New Para-ile Kn.m the Kulh'd (.nnisr. Jour. Parasitology 28 ill : 92. 205. J. S. H. 1886. [C(unments on Hunting Sea-on Prospects]. Forest and Stream 27 1 1 1 : 6. 206. Jacoiisiiagen, E. 1937. Millie unci Enddarin. In Bidk, Goepp<'it. Kalliiis iiml Luhosch, Handhuch der vergleichende analimiie der Wirheltiere. Berlin. 207. jAYtox, Thomas W., Jk. 1871. Bonasa Jobsii. Cornell Era 4 (12) : 182. 208. jEFEHiEs, George A. 1921. Raising the Rnffed Grouse. Game Breeder 19 l3, li : fiH. 1(17 lid. LITERATURE CITED 891 209. Johnson, E. L. 1924. Relation r,f Sheep to Climate. Jour. Agr. Research 29 (10) : 491-500. 210. Jones, Lynds. 1905. 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Field and Stieam 40 (3) : 73. 220. Lee, K. F. 1928. "Birds" and "Biddies". Forest and Stream 98 (12) : 748. 221. Lee, W. C. 1898. [Grouse Season in Manitoba.] American Field 49 ( 18) : 351. 222. Lehmann, Valcene W. 1941. Attwater"s Prairie Chicken, lis Life History and .Management. U. S. Dept. Interior, Fish and Wildlife Service, North American Fauna 57. 223. Leopold, Aldo. 1931. Game Methods — The American Way. .Am. Game 20 (2) : 20. 224. 1931. Report On a Game Survey of the North Central States. .Sporting .-Xmis and Animunilion Manufacturers' Inst., Madison. 225. 1933. Game Management. Chas. .Scribner's Sons, New York. 226. Leopold, A. and J. N. Bali. 1931. British and American (Jrouse Cvcles. Canadian Field Naturalist 45 (7» : 162-167. 227. Leslie, A. S. 1911. The Grouse in Health and l)i-ease. Smith, Elder & Co., l.ondoiL 228. Levine, p. p. 1932. A Report On an Epidenii( l)i-ease in Huff.-d (Iroiise. Trans. American Game ConL 19 : 437-441. 229. 1936. The Treatment of Ascariasis in Chickens. Cornell \et. 26 (2) : 120-127. 230. 1937. The \iahility of the Ova of Asraridia linriila W hen Exposed to \arious Environ- mental ('onditions. Jour. Parasitology 23 (41 : 368-375. 231. 19.38. The Effect of Infection With Davainea proglollina On the Weights of Growina Chickens. Jour. Parasitology 24 (6) : 550-551. 232. LiNGENKELTER, W. P. 1899. Game and Sliooting. American Field 52 (22) : 423. 233. LiNNAEiis, Carl Von. 1766. Systema Naturae 1 (2l 12th I'dilion. 234. Lloyd, Hoyes. 1927. Official (Canadian Record of Bird-Banding Returns. Canadian Field Naturalist 41 (7) : 161. 235. LoOMis, Leverette M. 1890. Observations on Some of the Summer Birds of the Mountain Portions of Pickens County, South Carolina. Auk 7(1) : 36. 236. Loveland, Clifton W. 1911. A Unique Experieme With the Ruffed Grouse Family Bird lore 13 (6) : 304. 237. Luttringer, L. A. 1935. Hawks- What They Eat. Nature .Magazine 26 (3) : 173175, 177. 238. MacFarlane, R. 1905. Mammals of the North-west Territory. Proc. U. S. Nat. Museum 28 (1405) : 673-764. 239. MacGrecor, Arthur E. et al. 1941. Summary of Grouse Crop and Gizzard Analyses for 1940. New Hampshire Fish and Game Dept. Tech. Circ. 9. 240. MacLulich, D. A. 1937. Fluctuations in the Ninnb.rs of the \arying Hare I Lepus americanus.) Uni\. of Toronto Studies, Biol. Series 43. 241. Macoun, John and James. 1909. Catalog of Canadian Birds. Gov. Printing Bur.. Ottawa. 242. MacVicar, Donald. 1918. Practical Grouse Preserving. Am. Game Prot. Assn. Bull. 7 (2) : 7-9. 243. Malthus. T. R. 1817. An Essay on the Principle of Population. J. Murray. London. 244. Marble, D. R. 1934. Relation of Juvenal Plumage to Growth and Sexual .Maturity. F'oullrv Science 13 : 195 201. 892 LITERATURE CITED 245. 1930. The Mi)ullin(! Faclur in ,liiilt;int; FhhU for Egg Prixluction. Cornell Uni%-. Agr. Exp. Sia. Bull. 503. 246. Marvin, C. F. 1929. Charactcrislics «i Cyrli:-. Krpnrls of Cimforences on Cyclis. Carni'gie Inst, of Washington. Supplemental Pnh. 1 : 11-12. 247. Massachusetts. 1900. Annual Report of Commissioners on Inland I'islieriis ami Game : 37-43. 248. 1906. Annual Report of Commissioners on Inland Fish and Game: 162-167. 249. 1936. Annual Report of the Division of Fisheries and Game : 16. 250. Mathews, F. Schuylek. 1921. Field Book of Wild Birds and Their Music. G. P. Putnam's Sons, New York. 251. May, J. B. 1935. The Hawks of North America. Nat. Assoc. Audubon Soc., New York. 252. Mayer, Alfred M. 1883. Sport With Gun and Rod in American Woods and Waters. The Century Co., New York. 253. Maynard, Charles J. 1881. The Birds of Eastern North America. C. J. Maynanl & Co., Newton- ville (Mass.). 254. McAtee, W. L. 1935. Food Habits of Common Hawks. U.S.D.A. Circ. 370. 255. 1936. The Malthusian Principle In Nature. Scientific Monthly 42 : 444-456. 256. McAtee, W. L. and F. E. L. Beal. 1924. Some Common (iame. Aquatic, and Rapacious Birds In Relation to Man. U.S.D.A. Farmers Bull. 497. 257. McDonald, D. 1935. Some Notes on the Habits of the Ruffed Giouse, Bonasa umbellus. Canadian Field Naturalist 49 (7) : 118. 258. McDowell, Robert. 1940. The Great Horned Owl. Penna. Game News 11 (8) : 10, 11, 29. 259. McIntosh, A. 1932. Some New Species of Trematode Worms of the Genus Leucochloridium Carus, Parasitic in Birds From Northern Michigan, With a Key and Notes on Other Species of the Genus. Jour. Parasitology 19 (1) : 32-53. 260. Mearns, Edgar A. 1901. Winter Birds of LakeWorden, Washington County, Rhode Island. Notes on Rhode Island Ornithology 2 (1) : 3. 261. Mershon, William B. 1923. Recollections of My Fifty Years Hunting and Fishing. The Stratford Co., Boston. 262. Michigan. 1936. Biennial Report, Dept. of Conservation (for 1935-36) : 203 206. 263. 1940. Biennial Report, Dept. of Conservation (fur 1939-40) : 241. 264. MiLLAis, J. G. 1909. Natural History of British Game Birds. Longmans, Green & Co., London. 265. Minnesota. 1916. Biennial Report, State Game and Fish Commissioner (.for period ending July 31, 1916) : 26. 266. 1924. Biennial Report, State Game and Fish Commissioner I for period ending June 30, 1924) : 37-38. 267. "Monon". 1875. Shooting Out of Season. Forest and Stream 5 (2) : 24. 268. MoRLEY, L. C. and P. W. Wetmore. 1936. The Etiology of Ulcerative Enteritis in Upland Game Birds. Science 84 (2177) : 272-273. 269. Morton, Thomas. 1637. New English Canaan. Prince Soc. Boston (1883). 270. Mueller, J. F. 1940. Parasitism and Disease in New Hampshire Ruffed Grouse Bonasa umbellus Lin- naeus. New Hampshire Fish and Game Dept., unpublished report. 271. 1941. Some Parasites Newly Recorded for the Ruffed Grouse, Bonasa umbellus, in the United States. Proc. Helm. Soc. of Washington 8 (1) : 14-15. 272. "Natura". 1887. A Swimming Ruffe of North American Bird>. Proc. I'. S. Nat. Museum 8 : 3.^4-356. 314. RiiM.KY. Ozark. 1939. Hawk Episodic. American Field 132 (33 1 : 171. 315. Rn'lM.K, R. v.. 1941. .Studies on the Gapeworni Synganius Irachrii i Monla^ii IKIII in Kohins and Chickens. Jour. Para.u-.- lla\<' Tli.ir "l p- an.l D.iwn-". Mi.liigau C.inserxalion 9 (12) : 2-10. 373. Tuttle, H. E. 1919. S.ime Notes i>n the Drumming .>f the Ruffe.l (;r..nse. Ank .36 (3) : 325-339. 374. Tyzzer, E. E. 1929. Coccidiosis in Gallinaceous Birds. Am. Jour. .)f Hygiene 10 (2) : 269-383. 375. 1930.. Flagellates From the Ruffed Grouse. \m. J..ur. .>f Hygiene 11 (1) : 56-72. 376. 1934. Studies on Histomoniasis or ""Blackhead" Infection In the Chicken and the Turkey. Proc. Am. Acad. Arts and Sci. 69 (51 : 189264. 377. U. S. Dept. of AcRin i.ti he. 19;?0. M..u>.- Cnlrol in Field and Orchard. Farm.rs Bull. 1397. 378. 19.36. Wildlife Handb...ik. I'. S. Forest .Service. R.-gion Nin.-. 379. 19.38. Outline .>f Procedure lor R.c.ir.ling Data Obtaine.l in St.Muach Exaniinatiiuis. Bureau Biol. .Survey Bi-1515. 380. Uttal, Leonard. 1941. Tarsal Feathering in Ruffed Grouse. Auk 38 dl : 7L 381. Van Roeckel, H. and M. K. Clarke. 1939. Equine Enceplial.imyelitis Virus (Eastern Type) Isolated from Ringneckeil Pheasant. Jour. Am. \el. Med. Assn. 94 (5) : 466-468. 382. Van Tyne, Josselyn. 1938. Check Li-t .>f Bir.ls .,f Mi.higan. O.ca-lonal Paper- 379. I'niv. of Michigan Museum of Zoid.igy. 896 LITERATURE CITED 383. Vreelano, Fiikdkrick K. 1918. How A Ruffed Grouse Drums. Am. Game Proi. Assn. Bull. 7 (1) : 1214. .384. Wmmo. W. W. and E. B. FoitnKs. 1941. Tli.- Chemical Composition of Forest Fruits and Nuts From Pennsylvania. Jour. Agri. Res. 62 (10) : 627-635. 385. Warren, B. H. 1890. Report on the Birds of Pennsylvania, Harrisburg. 386. Warren, D. C. and C. D. Gordon. 1935. The Sequence of Appearance, Molt and Replacement of the Juvenile Remiges of Some Domestic Birds. Jour. .'\gri. Res. 51 (5) : 459-470. 387. Wayne, Artiii-r T. 1910. Birds of South Carolina. Daggett Print. Co.. Charleston. 388. Weed, C. M. and Ned Dearborn. 1903. Birds in Their Relation to Man. J. B. Lippincott Co., Phila. 389. Wehb, E. E. 1926. Earthworms as Transmitters of Capillaria annulala. the "Crop-Worm" of Chickens. N. Am. Vet. 17 (8) : 8-20. 390. 1937. Observations on the De\elcipnient of the Puultrv Gapeworm Syngamus trachea. Trans. Am. Mirr. Soc. 56 H) : 72-78. 391. 1940. A New Intestinal Roundworm From the Ruffed Grouse in the United States. Jour, of Parasitology 26 (5) : 373-375. 392. Werby, H. J. 1928. The Trematode Genus Hormostomum. Trans. Am. Micr. Soc. 47 (1) : 68-81. .393. Wetmore, Alexander. 1937. Observations on the Birds of West Virginia. Proc. U. S. Nat. Museum 84 (3021) : 401-441. 394 1940. A Check-List of the Fossil Birds of North America. .Smithsonian Misc. Coll. 99 (4). 395. White, F. B. 1929. Birds and Motor Cars. Auk 46 (3) : 399. 3%. Whitehead, F. E. 1934. The Effect of Arsenic, as Used in Poisoning Grasshoppers, Upon Birds. Okla. Agri. and Mech. Coll. Agr. Exp. Sta. Bull. 218. 397. Wight, H. M. 1930. Pheasant Management Studies In Michigan. Trans. .Am. Game Conf. 17 : 222. 398. Wiioi s, H. S., Jr., L. C. Norris and G. F. Hedser. 1937. The Role of Manganese and Certain Other Trace Elements In the Prevention of Perosis. Jour, of Nutr. 14 : 15.i-167. 399. WiLi.ETT, George. 1914. Birds of Sitka and Vicinity. Condor 16 <2i : 89. 400. Wilson, Alexandeh. 1812. .Vmerican Ornithology. Bradford & Inskeep. Phila. 401. Wing, L. W. 19.34, Cycles of Migration. Wil-an Bull. 46 (3) : 150-1.S6. 402. 1935. Wildlife Cycles in Relation to the Sun. Trans. Am. Game ('.«»(. 21 : 345-363. 403. 1937. Wildlife and Cycles. American Forests 43 (lOt : 512-513. 404. ViouuRiKF, E. Seymour. 1908. The Ruffed (Arouse, A Study of the Causes For Its Scarcity in 1907. Annual Rept., N. Y. State Forest. Fish and Game Commission, 'for 19071 : 1,59-178. ■U)5. Yapp, R. H. 1922, The Concept of Habitat. Jour, Ecology 10 : 1, INDEX 897 INDEX nt of 597 Abbott, C. C. 3,260 abandoned lands, importance and cncouragen abundance, grouse (see also fluctuations) early records 2, 6, 11, 53, 558. 560. 562 ' factors governing (see also factors) 7 reduction of 557 relative 53. 55. 557, 559, 676 Acarina 409, 775 accidents (see mccbanical injury) Accipiter cooperi 317, 333 Accipiter vetox 317, 333 Acer 198. 204 Acer pennsylvanicum 204, 8 18 Acer rubrum 204. 848 Acer saccharum 204. 848 Acer spicalum 204 Aeeraceae 204 Ackerl. J. E. 424. 426 aconitic acid 195 acnrn 188, 202. 215. 221 action, sex differences 44. 45. 165. 167. 840, 843 Adaina, C. C. 32 Adams. W. C. xxxv adaptability to changing environment 272 Addy, C. E. 31 Adirondack Forest Preserve (N.Y.) 596. 684 Adirondack Mountains (N.Y.) 3. 36, 54, 56. 57. 95. .384. 386 Adirondack region (N.Y.) absence o( stomach worm records from 415 characteristics of 114. 217. 2M. 596. 694 cover management in 596 Adirondack study area (N.Y.). description 695, 696 adult cover basic patterns 593 composition 593 density 593 m on t illy and seasonal preference.'* l.*>3. 819 relation to roosting habits 163. 168. 836 relation to slope 594 relation to time of day 159. 823 relation In water 594 relation to weather 161. 831 relative importance u( Ivpea I.'il types choaen 152, 156. 819 undergrowth 594 use during hunting season 160. 829 adult mortality (see also survival during adult period I 31B cliief predatftrs involved (see also methods aad teohniqiiefi) 308. 322. 337, 348, 669 effert of predator control 308. 328. 347 number of dead birds cxaminetl 336, 532. 876 proportion attributed irt \arious causes 532 (jiuilily of eviileure ascribed to tiiai:nosi< u\ ilead birds ex- amined 876 recorded during Invesligalion :U)8. 3 IK iclation to accidents 318. 531 iclation to disease 319. 332. 531 lelalion to fall population density 308. 320. .331. 338. 532. .133. 534. 535. 552 relation to maturing chicks in fall population 535. 536. 5U relation tfi variations in buffer abundance 322, 532, 535, 513. 546. 552 relation to variations in predator pressure 321, 535, 552 relation to weather 299. 304. 321. 332. 532. 535 role of predalion 308. 319, 322. 332, 333. 317. 531, 532. 535 seasonal variation 322, 336, 338, 5.36. 537 iiftcrshaft 59, 747 Agamodistomum 409, 434 age breeding 359 how to judge 84. 718 longevity 360 iclation to feather devclupniciil 37. 81 age composition of populations 513 relation to adult survival 535. 536. 537. 545 significance in Huctualions in abundance 571, 573 agricultural productivity, cycle in 578 agriculture (see farming) Agrimonia 193 airplane, use in transporting eggs 152 air-sac mile 409. 415. 417. 436 air-sac mite infection cause 441 control 441 tlissemination 413. 441 nature 441 .'Vlabama 236 Alaska 36. 48. 52. 53. 54. 110. 236. 285. 325. 409. 128. 434 Albany County (N.Y.) 199. 213, 433. 520. 695 Alberta 18. 20. 47, 48, 53 albinitiU) 57. 58 albumen 74 alder 212. 235. 244 alder beds, importance in Adirondack region 140 Aldrich. J. W. 48 Alger County (Mich.) 847 alimentary tract (see anatomy) alkaloids, toxic 195 allantnis 74. 76 Allegany County (N.Y. I .568 Allegany Mountains 53. 217 Allegany State Park (N.Y.) 684 Allegheny National Forest (Pa.) 312 Allen. A. A. xxxi. xxxvi. 15. 22. 24. 27. 29. 30. 32. 58. 1^3. 187, 210. 2.56. 267. 274. 275. 278. 282, 284. 288, .333. 415. 419 421. 447. 453. 454. 457, 462. 476. 508. 568. 574, 694 Allen. G. M. xxxvi. 721 Alnus 212 Alopex lagopu-s 323. 578 Ambrosia artemisiifolia 206, 819 Amelnnchier 198. 200 Amelnnchier canadensis 817 Amelnnchier laevis 847 American Game Conference 12. 29, 32. 33 American Game Protective Association xxxvi. 12, 22, 25. 29. 30, 153. 562 amnicm 74. 76 imphicarpa monoica 8-18 Amsterdam (N.Y.) .371 amygdalin 195 .\nacanliaceae 203 anatomy 58, 721 alimentary tract (see digestive tract* comparison of various genera of Tctraonidac 58, 724, 725, 726, 727. 728 fligcstive tract 58, 737 distinctive characteristics 58 movement r>f leg 734 musculature of leg 727. 729 musculature of wing 735 respiratory system 58 • v^^ ^i skeletal measurements 727 ,» ^ A skeleton 721 "^ "^ structure of leg 726 structure of pelvic girdle and ••ynsocFum 726 structure of rib-i 725 ^ r-truclure nf skull 721 * ' -iriiclurc of «ieniuni and pectoral girdle "25 «truclure of »iyrinx 58, 737 * structure i»l vertebral column 721 -tructure of wing 725 uro-genilal system 58. 734 andromedotoxin 193. 195 anemone 208 Anemonr 208 ant 101. 182. 188. I'».V 198. 212. 213. 221. 221. 22.5. 261. 408. 776 ani. as a source of lapetturm infection 426 ant. carpenter 213. 218. 819 anI. cornfield 213 ant. wor»d 213 ant family, a* a source of grouse food 212. 221 Antennan.i 198. 201 Antiosti Island (Que.) 18. 20. 50. 254. 368. 501. 508 aphids 776 ipios tuht'tosa 818 -\|>palacliian Mountain' apple 188. 198. 199. 205, 225. 226, 227. 228. 231, 6.55. 662, 707. 8^17 apple-tree worm 101, 212 Aproctella stoddardi 409 Aquiia chrysaetos 333 Arachnida 198. 213 Araneida 775. 776 Arctium 193 Arrtium minus 819 Vrkansa^. 46. 50. 52. 272 Armadiliidium tulgare 421 army worm 101 ironia 200 arrangement of cover (see interspersion ) arrowwood 203. 204. 655. 662. 849. 883 artificial propagation («ce also methods J 691 biological limitations 416. 499 comparison of quail ami grouse 158 cost 502 difficulties 416. 500 diseases encountered 446. 118. 157. 190 early attempts 21. 447. 453 effect of breeding pen arrangenwrnt on egg productio effect of brooding conditions on chick survival 881 effect iiicti1 21. 147, -i5:t birfcli. Kurupeaii 210 fxpcrinifiitH (iindurtvt] at Cati*kill fi«*M slalion H8 heec)) tainily. as u sniinc of groubi- foiM 201 (.-xptTimenls rontliictctl nt Krsrarrli Center 877 Iieerhdroii 210. 218 rartors liniitinK proaiH-lum U6 500 h.-i-rhiuit 188. 'l89. 197. 202. 201. 215. 221. 2;W. 2i;( fertility of ef-ps (r<»in birds lieM on groiin.l 151 flmluatioii in . r..t. '0' ferlilicv nt ecK* from birds held ..n wire 470 .piantili.-. . on.i.M.r.r 201 food rations fur reaniit; younp pronse 187. 2.'{7. 119. I.i(». l«K. HHli jj^.^.^. j ^^ .,Q■^ Ills.;;!, ;';'^ .., R,-,o„r..i, c,-.,..t bts I'-'i- j^yf- •««■ "?„,^'?;,--'- --'•--='• -^'- '"« in,ubat„r.bru,„l,.r ll,...l 157 '>■'■"■• ' '"'' ''\" , ,'"'■ -'- lack of traini-,1 ,„a,„,„w.., 501 '"■^''^•- ••,"'■",'"';", n','^'.;,-,'- liberarion „r hir.ls r.-ar „■>■ lil.,„.,i..„ I beel e. ol,„ l.-at 101. 212 matine bi-havio, 471 "■■•"■ f"r„,„-ai, .-In, U-a( 21.i mclho.U I see melh<..lsl '"■"■ "-""I"' ;"";,!;■"',"" natural rover melho.l 150. 1.5.i |"-''l!<-- f'",'"".'....-'..'.-. "' obtaining bre<^-din(: Hio<-k .501 prarticability 116, 1.50. 197 prcdation 4.56. 490 tirodiiotion eqnations 498 rearine .urres. at Researeb C-n.,., I Asia 426 as|ieet (see slope! aspen 115. 182. 19.5. 198. 199. 201. 211, 214, 216. 217. 218. 21-,, „ , , , ,„, 220. 225. 226. 228. 231. 2,i2, 235, 238. 239. 24.-). MO. 707 ""■'""'"""':/'"• -°' 1 . .1 I .in I .11- .1..- Metillareae 2W) aspen, lariie-tuothed 201. 21.1, 22i R^,,!,.!, H U aspen, tremblinu 186. 189. 201. 205. 207. 211. 21.5. 227. 2.57. 211. , - , V i- , ' , „, " big bnek disease {see perosi?.! -u .11 11^ ,1- ,» Bie Horn Mountains (Wvu. and M„ afiiercil osis 111.416. 11 .. 1.16 „.\,. .. , „,- ' *^ ,~Q Rilbngs. K. 6. 260 ""T I in 10/; Binphanilnn ).N.Y.» 254. :t9I eontrol 441. 496 i - i ■ i i t i .t-,^ ,. .. ,., biologu-al assuv of food 2.i6 di9»eminatinn 141 u- i ■ , i i " ■ ,--„ ,..-, binlopicat balanee slieel .550 nature 4.i9 , . '^. . , , . , , beet).-, b-af 212. 221 beetle. \lii\ 101. 21,1 beetle, pair slri]M-il flea 101 beetle, searab 224 beetles, af^ ti smiree of crrtiise fmid 21it behavior, display of dnniinanee .17. 61. 268. 2HI beba%ior. mating ;{7. ftli. 65 Herpen (N.Y.I Hergtold. W. H. .50 fl.-ruhi 198. 200 Hflulit Ifnln 201 Herula lutra 200. 816 , n t ■ . , 1 11 ■ I ii.o <><> bioinelriral analvsis of data 110. .:>.io. 510. 701. 70H. 71H A.,.;nw.ll T 1-1 anal>8i8 ol \arian.e -19 .-Xspinwall. T. 14 ehi Ill-nil lini I- 1 1 :r "*!'I'^'"->ni •>>o repression and eorrelation 535. 708. 720 4 / "ofti " sipniliranee 719 . t 1 ( . I .... l''^*-!' 56. 115. 182. 198. 200. 215. 216. 217. 218. 219. 220, a*ter family, as a soiiree of pronse lood 201 .,.,^ .,,. ,,„- _^, ' ... 1 1 i »i. e 1 . cr bireh, blark 201. 2l3. 225. 22<. 235. 6.i.t Atbaba»ea Lake (Alta., ^ask.) 56 k- i . . , ,... , ., , birch, caniM- (see paper I atinoApheru- ronilitions isee weather! i- . .,.,. , ,1 A,„l„bon. J.J. 6. 14. .52. .53. 2.53. 2.5.5. 271, 276, 333, 371, 3»i. I""',''- f"" "^^^]° •Vii Sable Slate Forest (Mieli,! 817 aiiloiiinbile (see man's aids) avens 193, 200 A>is, W, H, 14 bireb. pa|><.r 201. 215. 227. 610 bireh. white (see paper) bireh. yellow 200, 205, 209, 211. 213, 225, 227, 237, 210. 244,26,5, 610. 655. 816. 88.3 . ... . e — variation in rlieinieal eoinposition of t>ii«s 2 tO, 8 6 Awenie (Man,) 5.vi i ■ i r -i t r i ..^,. , • ,1 . birrii fainilv, as a soiiree of croiise fooil 200 ■*.■ (see inuil s auls) ■ - am lea 203 bird. pox , 208 Hiseboll ,52 llislio|.. J, S, ,11 I'abrork, C, 14 bishop's eap 206 Kaekus. E. E. 22. 24 bittersweet 190, 191, 195, 635. 662. BRl barleria 239. 408. 668. 671 blaekberrv 182. 198. 199, 215. 216. 218. 219. 220. 221. 222. 223. baj; limits (see hunting re|:iilations) 224. 225. 227. 231. 243, 655 fiailey, H, H, 48 bla.kberrv , Bailey's 847 Hailey. tt, « . 183 blaekbird 196 Baird. S, F. 47. .52 blarkeoek 2. I. 25 Bill. J.N. 3D l.la.klly 108.433 balsam fir 56 blarkbaw 883 Balsaminaeeae 208 blarkhead 402. 104. 116. 133. 136 llanits. (). 18. 47, 48 eaiiae 128, 138 Hanta, f,. 23. 30 eontrol 138. 150. 172. 192 banlani hen. use in real inn trolls,. "7, 157, 171 disseniinalion 138.4.53 Itarbi.r. If. (^. xxxvi nature l.tR ll,itnliard, W. 18.19 lllaek Hills ( So. Da. and Wvo. I 30, ,133 llarllett, M. T. 102 lllaek.burK (Va.l 202 llarttaln. J. 3.6.7.13,21,16,271,276 blaeksnake .133. ,33 1, 876 • Knrlseh. V. xxx\\ Blair. F. I). .577 l!a... C. C. 436 bli|:hl. eliestniit 231. 232 II lie. J.R. 2.3 blood iiara.ite. Baten. W, I). 760 ileserijplion 130 bavberry 8a3 distribution 130 Bial. F. F, I„ 10) no i.lenee and iinporlaiiee 1.30 bean, wild 848 life rvrle and ,liss<.|iiinaIioii 130 Beaudelte. F. K. 113 lutliouenieitv I3U beaver 683 blue.beeeh (sei, loirnbeain. aineriean I Beek. T. 2,3.278 blueberry 189. 190. 202. 220. 222. 221. 225. 231. 232. 655. 819 be,lslraw 208 blueberry, low liiisli 227 1 223, 221, 309. 776 bluebird 431 beeeli 11.5, 198, 201. 202. 209. 215. 216. 217, 218, 219, 220. 225, bobeal .330. 3,33, ,131. 382, 711, 876 226, 227, -230, 2.12, 610, 6.53, 707. 816, 88:1 b,.l.wbile uaill INDEX 899 265. 274, 276. 279. 572 boJv lemperaiure (see also physiological studies) 62 Bonasa 46. 724. 725, 726. 727, 728, 730. 732, 733, 737 Honasa ceres '16 Bonasa, characters of 46 Bonasa jobsii 48 Bonasa umbellus brunnescens 48 Bonasa umbellus helmei 48 Bonasa umbellus sabini 47. 48. 52. 53, 72, 235 Bonasa umbellus ihayeri 47, 48 Bonasa umbellus togata 39. 47, 48. 53. 261 Bonasa umbellus umbelloides 47. 48, 53, 197, 210 Bonasa umbellus umbellus 38. 39, 47, 48. 53 Bonasa umbellus yukonensis 47. 48. 197 Bonasia major Canadensis 47 bonasus 46 borer, poplar 101, 212 Boston (Mass.) 8. 557 botulism 671 Boudoin College 33 Boughton. R. V. 422. 42.H bounties, on grouse 11, 37. 102, 187 bounties, on predators 381, 382, 383, 388 Rrarhyrhinus rugifrons 213 bracken 309 bracken. E. A. 23 Bracketr. M. W. xxxv Bradbury. H. M. 243 Bradley, J. T. 360 Bradshaw. F. 15 Brazil 419 Breckenridge, W. J. 521 breeding (sec also mating liabita) readiness for 69. 267 relation to age 353, 359 breeding failure (see also non-breeding) 311, 353. 355. 539, S breeding season 265 breeding stock, sale of 104 breeding success (sec productivity) Brewer. T. M. 52 Brewster, W. 255. 260. 261. 262. 264. Bridges. H. P. 23 Brisson. M. J. 47 British Columbia 31, 47. 50. 52. 53. 51. 193. 409. 435 British Columbia Provincial Museum 31 British Committee of Inquiry on Grouse Dificase 40t British Isles 103. 104, 578 Bronx County (N.Y.) 56 brood cover 138 basic patlerns 589 composition 593 density 593 efTecl of density of crown cover eflerl of density of undergrowlh influence of slope 148, 593. 813 influences affecting choice 138, 801 relation to age of brood 141 relatifin to ground conditions 141, 8117 relation to lime of day 142, 803 relation to water 593 relation to weather 142. 147, 305. 801. 810. 815 relative importance of types 139. 801 types (if crown cover chosen 139. 801 types of undergrowth chosen 146, 808 undergrowth 593 brood mortality (see also survival during brood period I 3 causes responsible 528 chief predators involved (sec also methods and lechniqu*-' 335, 385 fomparison with artifically reared stock 316. 528 cfTeri of predator contrnl .^08. 316. 347 number of dead birds examined 335 i)tialilv I'f pvidciiri- ascribed to dlagmiiies of dead bird ined 876 recorded during Investigation 315 relation to weather 299. 303. 317. 528 role of predaiion 307, 317. 347. 528 brood spot 59. 746 broods (see also chicks) area traversed 248. 291 double 293 habits of 294 Brooke, Mr. 13. 274. 276 broomrape family 210 Brown Sand Hills (Tex. I 52 Briinelt. E. L. 33 Brunswick (Me.) 34 Bubo virginianus 322, 333 buckbrush 203. 204 buckwheat 210, 212. 230, 233 buckwheat, common 847 buckwheat, false 847 buckwheat family, as s source of grouse food 206 budding 37, 103. 187 141. 802 147, 800 buffers, (see also methods of management) fluctuations in abundance (see also methods and techniques) 308. 322, 325. 556, 567 principal species of the Northeast 325 relation of fluctuations to fox activity 308, 322 relation of fluctuations to predation 307. 308. 315. 322, 552 bug 213. 224 bug, plant 101. 212 bugs, true 213 building activity, cycle in 578 Bull Hill (N.Y.) 345. 520 Bump. C. 1. 15, 22. 24. 30. 34, 35, 63, 90, 100. 105. 181. 183. 220. 443, 550. 557, 568. 581, 585. 605, 637. 667. 674, 694, 845. 846, 847, 848. 849 Bump. J. 22. 24, 30. 694 bunchbcrry (see dogwood) Burckmyer, E. K. xxxvi. 33 burdock 193. 849 Burnham. J. B. xxxi. xxxvi. 15. 27. 30. 32. 95, 264. 562, 574 bursa Fabricii 84, 90, 94. 256. 434. 516. 718, 739 Buraon, E. H. 14 Buteo 323, 330 Buteo borealis 330. 333 Buteo lagopus s. johonnis 330 Buteo lineatus 330. .3.33 Buteo platypterus 330, 333 buttercup 208. 218. 222. 223. 224. 231 buttercup, tall 847 buttercup family, as a source of grouse food 208 butterfly 213 calcium, o food component 239 calcium oxalate 195 California 2. 36, 46. 272. 433, 671 calls 99 Camden (So. Car.) 52 Camponotus herculeanus 213, 849 Canachites canadensis 210 Canada 4. 8. 18. 30, 36, 48. 52. 53. 54. 56. 104. 241. 323. 325. 396. 505. 557. 562. 563. 565. 567. 568. 572. 579 Canandaigua (N.Y.) 208 Canis 333. 578 canker-worm 101, 212 Cape Cod (Mass.) 453 Cape Mendocino (Colif.) 47 capercailzie 25, 427, 434 Capillaria 402, 408 Capillaria annulata (sec also cropworm) 409, 416. 419. 425 Capitol District Came Bcfuge (N.Y.) 18 Capri folioceae 204 Carabidoc 213 carbohydrate, a food component 236. 238. 239, 240 Caret 198. 203 Carex crinita 203 Carex debilis tar. rudgci 203. 222 Carex gracUlima 222 Carex intumescens 203, 222, 845 Carex lurida 845 Carex viridula 222 Cariboo Mountains (Br. Col.) 53 Carnegie Institute of Washington 25 Carpenter. F. H. 558 Carpenter. R. C 102 Carpinus 198. 201 carrot, wild 193. 848 carrying capacity 309, 332. 393. 512, 522. 546. 672. 677 criteria 523 definition 523 observations of others 524 values recorded by Investigation 523 variability 524 Carticr. J. 46 Cartwright. B. W. xxxvi Cascade Range 47 rat, house 14. 25, 330. 333. 334, 336. 338. 339. 340. 311. 34^1, 385, 391. 456. 496. 711.875. 876 catbird 420 caterpillar 101, 182. 195, 212, 224. 231 caterpillar, forest-tent 232 Catskill Experiment Station (N.Y.) 297. 301, 4-t8, 4.i5, 475. 465. 492. 502, 504 Catskill Forest Preserve (N.Y.) 596. 684 Catskill Mountains (N.Y.) 56. 57. 115, 386. 4^18. 599 Catskill region (N.Y.) characteristics of 115. 217. 596. 694 cover management in 596 Catskill study area (N.Y.), description 695 cattle (see livestock) Caulophyllum thalictroides 195 causes of fluctuations in grouse abundance (see also productivity) 571 general 511. 571. 573 relation to age composition of population 573 relation to disease 309. 401, 574. 575 relation to population density 556, 576, 577 900 l\DFX relation to predalion 574 relation to solar activity 575 relation to weather 556. 575, 576. 577 role of adult mortality 548 signi&cancc of success or failure of annual increment of young 534. 556. 571. 572. 573, 577 Caulheo. G. E. 42* cave. Cumberland (Md.) 2. 46. 272 cave, Frankstown (Pa.) 1. 46. 272 rave. Potter Creek (Calif.) 2, 46. 272 Cayuga County (N.Y.) 19 reral worms description 415, 425. 427 distributinn '409. 416. 427 incidence and importance 410. 428 life cycle and dissemination 408, 428 pathogenicity 428 cedar, white 655. 660, 883 Cedar Lake (Tex.) 52 Celastrus scandens 194 cellulose, a food component 239 census of grouse populations (sec metiioda and techniques) Centre County (Pa.) 846 Cen/rocerco5 724, 725 Ceratophyllus diQinis 409, 435 Cestoda 409 Chaddock, T. T. 183. 208. 212, 220 Chamberlain. W. J. xxxvi Chapman. F. B. 30 Chapman. F. M. 38 characteristics (see anatomy, taxonomy) Chateaugay (N.Y.) 202 Cheilospirum 408. 494 Cheilospirura hamulosa 423 Cheilospirura spinosa (see also gizzard worm) 106, 109, 413, 116. 423. 425 chemical composition of grouse foods 845 chemical poison, relation to disease 404. 405 Chemung River (N.Y.) 57 Chenango County (N.Y.) 10. 95, 222. 407, 520. 558. 695 Chenango public shooting ground (N.Y.) 391 Chenango vallfV (N.Y.) 695 cherrv 182. 186. 188. 195. 198. 199. 214. 215. 216. 218. 219. 220. 221', 223. 224. 225. 231. 235, 237. 238, 239, 24-1. ,595, 707 cherrv choke 199. 207. 211. 215. 226. 227. 2.30, 847, 848 cherry, pin ll.V 190. 199. 209, 215. 221, 222, 225, 226, 227, 228. 232. 610. O.Sr.. 817 cherry, wild black 199. 205. 215. 225, 227, 228, 610, 655. 817. 883 chestnut 188. 202 Chicago (Ml.) 8 chicken (see fowl) chicks 248 ability to hide 295 calls 99 dust bathing 297 feeding habits 221. 296 grcgariousness 297 growth and development 78 roosting 296 Childs and Walcott Preserve (Conn.) 27 Chilopoda 775 chipmunk 28, 333. 334. 335. 340, 341. 670. 711. 876 Chippewa National Forest (Minn.) 521 chi-squarc (sec biometrical analysis) chitin. a food component 239 rhloro[)hyll. affericd by solarization 242 Choanoliivnia infundtbulum 409, 434 chokeberrv 200 Chrysomrlidac 213 Cincinnati (O.) 8, 371 Circus hudsonius 330, 333 Civil War 557 Civilian Conservation Corps xxxvi dam. finger-nail 192 Clare (Mich.) 19 Clark. G. W. 47 Clark. M.W. 21.22.24.453 Clark. T. E. 31. 183 Ciurk University 23 Clarke. C. H.D. xxxvi, 31, 52. 56, 312. .324. 430. 433. 55*. o60, 562. .566. .';67. 568. 572, 574 ctassifiration 46 clear-cut areas (see slashings) clearing of land 383 relati4 occurrence 6. 7, 11. 12. .5.55. .5.56. .571 periodicity 555. 561. .571 rate of change 555, 5(>0 relation to behavior of local populations 555. 568, 571. 577 relation to population density (sec productivity, net) sequence between regions 555. 567 synchronism 555, 565. 571 years of principal declines (1900-44) 566 flukes 409. 434, 671 flushing distance (see distance grouse flush from observer) fly larvae, raising 456 foamflower 206. 220, 227 FoUeit, R. E. 19. 23, 24 food conditions, recording 698 food habits of grouse (see also feeding habitsl 181 competition with other speiics 230 materials studied 184 methods of study 182. 184. 706 relation to other faclois of abundance 241. 511, 513 seasonal differences 182. L'l 1 studies of 30. 31. 183 food of grouse amount eaten 189 chemical composition 237. 845 choice in relation to nutritive value 238, 240 effect of opening up the woodland 231 effect on grouse distribiilion (local) 182, 229 general availability 182. 227. 229. 528. 775 items often considcied injurious 193 miscellaneous items 188. 192 monthly variations in food of chicks 224 nutritive value 237 of young grouse 182. 221. 415 other vertebrates taken 188. 192 principal chick foods 221 regional differences in New York 217, 227 relation of food composition to solar activity 242 relation of food composition to weather 242 relation to cycles 241 relation to health of the bird 235 relation to weight of the bird 37. 241 seasonal differences 182. 214. 227 types eaten 197 variation throughout range of grouse 182. 220 variations in composition 237 variety of choice 182. 191. 198 yearly differences 182. 218 food plants distribution in New York 227 effect of lumbering on 233 habitat requirements 225, 227 relatit frequently taken 198 variety eaten 198 yearlv differences in bulk taken of important species 218 Fnoie. I.. 562 Forbes. E. B. 202. 816. 847. 848. 819 Forbes. S. A. 194 Forbush. E. H. xxxi. ID. 12. 13. 14. 30. 38. 98, 102. 252. 2.56. 260. 262. 264. 269. 270. 292. 333. 3,38. 371. 375. 561. 572 Fordham, S. 22. 24. .30 forest, climax .53. 112. 386. .393. fi94 forest, sub-climax .53. 51 forest districts 687. 688 forest management for grouse 387. 587, 599 Forest Practice Ad .598 forest preserve (sec also Adirondack; Calskill) 684 forest products 587 forest BtantI and game cover improvement 6.50, 777 forest succession 118. 386 forestry, control of insects and disea<>i-s 654 forests, national 682 forests. State 682. ()83 formic acid 195 Formica 213 Formicidae 213 Fort Reliance (N. W. Terr.) 52. 56 Fortymile (Y. T.) 47 fossil records 36. 46. 272 fowl, domestic 239. 242. 408, 412. H5. 116. 118. 419. 420. 423. 424. 426. 427, 428, 430, 434 as a carrier of disease 385. 415. 417. 434. 435. 138. 439. 411 association with grouse 268. 290. 385 fowl, guinea 420. 426 fowl cholera 404. 415 Fox. .K. xxxvi fox (see also various species) 14, 313, 321. 323. 329. 330. 339. 350. 391. 417. 552. 630. 667. 669 as a predator during adult period 308. .122, 337, .138. 348. 876 as a predator during nest period 307. 308. 314, 334. 347, 527. 669. 710. 711. 876 as a predator of grouse chicks 335. 336. 876 fluctuations in abundance .328. .567. 579 food habits 308. 327. .3,19. 340. 341, .142, 344. 709 relation of variations in fox activity to adult mortality 322 relation of variations in fox activity to buffer abundance 308. 322. 328. 670. 708 relation of variuiiims in fox activity to nest mortality 315 fox. arctic .123. 578 fox. grav 314. ,324. 329. 332. 3,36. .137. 339. 381. 382. 683. 711. 875 fox. red 314. 323. 324. 329, 332, 336. 337. .139, 381, 382, 578. 683. 711. 875 Foxborough (Mass.) 10.375 Fragaria 186. 198. 199 Fragaria virginiana 847 France 3 Franklin County (N.Y.) 19.558 904 INDEX Frascr River (Br. Col.) S3 Fraxinus 212 Frederick. K. T. xxxvi Fredericks. W. 24 Fredine. G. 560 .Frie and uann- nianagrmeni 623 interbreeding 268 inlerplanling 660 interpredalor relalion>-hips .'III inlerspersion. role of 113. H>8, 609. 818 intestinal inflammation 411. 412 intestinal worms (see also large roundworms) 15. 108. 409. 410. 411. 412. 413. 416. 125 intimidation display 66. 282 investigations reganling grouse xxxi. xxxii. 30 I own .10. .50. 183, 188 Iowa .Agricultural Experiment Sialiun 30 iron, pig 578 ironwood (see hop-hornbeam I Isham. R. H. 23 Ishii. N. 31 Isle Royal (Mich.) 50. 251 Isopoda 775 Ithaca (N.Y.) 183. 199. 203. 213. 239. 255, 302, 325. 579, 747. 815. 816. 817. 818. 849 Ithaca Came Farm IN.Y.) 18 ivy. grounil 195 ivy. poison 194. 195. 203 Jackson. H. H. 15 Jacobson. K. .\. 31 .Kicckel. H. F.. Jr. 21 Jay Mountain (N.Y.) 695 JefTerson Countv (N.Y.) 199. 213 Jeflries. G. A. 16. 22. 24. 27. 30 jewel-weed 207. 208. 215. 218. 222. 221 iewel-weed. pale 215 jewel-weed, spotted 207. 215 irwel-weed family, as a sotirrr of grouse food 208 ]..!., H. K. 22. 28 906 INDEX Johnson, R. A. 30 Jones. J. C. 181. 183. 220, 339. 694. 845. 846. B4T. 818, 819 Jont^s. I.. 276 Sunn. M. C. xixvi J..n.-». M. F. 427 Jun (N,Y,) 8 Kinnev, A. B. F. 14 KniEht. O. W, 72 Kiihn. T. M. 183, 220 Kuser. A. R. 23 U Pierre House (N. W. Terr.) 52. 56 Labrador 47. 56. 409. 415, 428, 134 I.atli], C. E. xxx%'. xxxvi l.ae)ap(jnae 409. 417. 434 I.agopofcus perptexus 409, \\\. 135 I.agopus 724. 725. 727 I.iignpus lagopus 578 I.agnpus scoticus 578 Lahonton, Baron de 3. 100. 260. 274. 276 Iviing. H. M. 15 Lake Chaniplain (N.Y. and Vl.) 3. 326. 345. 415 Lake Superior 50. 254 Lake UmbaeoK (Me.) 25,5, 2tl himl clearinp (see rlrarinK of land) Lantz, O.K. 341 lapwinc 434 larrh 115 lareli, European 655 l.arix laririna 815 l.asius niger 213 laur.I 193. 195. 198, 203. 205. 210, 218. 235 latirel. inniinlain 186. 189, 193. 203. 215. 218. 227, 2.37, 238. 214. 849 laurel, sheep 203 laurel puisnninp 193 Lawrence, W. M. 694 law8 (see ronserration ; hunting regulations) Lawyer. C. A. xxxi. \xxv. 34 lead arsenale 196 lead poisoninc 15. 192, 10,5, 411 leafhopper 101. 212. 776 LeComple. E. L. 23, 24 LeesMrRae Collece 23, 24 I.effinBwell. n. J. 30. 33 Ice (see ana(urny) Legge, L, xixv lepume family, as a souree of grouse food 208 I.egiiminosae 208 Lehman, R. 19 Lehmann. V. ^'. 243 li-mmiiiK (nee also buffers) .323. 325. 556. 578. 579 I.emmus Irmmu* 325. 556 t.emmus ttimucronalus 323, 578 Leopold, A. xxx\\. 12. 30. 266. 310. 326. .367, 375. 390, ,5,56. .557. .558. 559. 560. 562. .WiS. 568. 574. 575. 577. 674 Lepidoptera 198. 213. 223. 224. 775 I.epu-s iimrriranui 322, 5.56 l.fucochloiidium pricei 409. 434 Lrucocylozoon 430. 571 Leucocytozoon anatii 433 t.eucocYtozonn honnsar (see also hlood parasites) 409 433 Levlne. P. P. 101, 424. 437. 495. 694 Lewis, H. F. xxxvi Lewi*. M. 47 Lewis Countv (N.Y.) 21 L 4.33 Liard Cap (Bt. Col. and Y.T.) 48 Libbey. n. S. 14 liberation of artifii-ially reared grousr farlors nITerline survival ,506 feeding .504. 505 marking .505. 716 method of lihernlton 501. 673 survival rernrded 18, 508 Iranspnrlaiion 50-1 liberations of grousr, records of 18. 19. 150. ,507 library American Museum of Natural Mislorv xxxvj Cornell L'niver«ity xxxvi New York City xxxvi New York Slate xxxvi lice (sec louse I life equations of grouse 526. 546 basic differenrcs recorded between stable, increasing and de- creasing abundance 516, 547, 518. ,519 effect of sport hunting 538. 516. 550 lignin, a fit' 19(» Mainntbrmum ranadensf 210. 845 Maine 8, 22. 24. 31. 72. 235. 255. 261. 396. 409. 424. 1.33. 435. 558. .563. .566. 567 * malaria-like parasites (sec also blood parasites) 408. 409. 430 male grouse (see also courtship: drumming) habits 271 relation to broitd 218. 284 relation m nest 218. 284 Mahhu-.. T. K. 512 \faltluisiaii principle 51.1 }t,iiu.s 198. 199 Vtiliis piimi/a 847 man. as inlluence on grouse abundance 369. 527 as a conservationist 371. 387 as a farmer 116. 370. 371. .383. 597 as a grouse hunter 370. .371 as a hunter and trapper of predators 370. 380 as a lumberman 233, .370. .371. .385. .597 studies of .30 management ("ee also methods of management) ad%ice on 652 designing grouse coverts 607 effect of Forest Preserve amendment on grouse rover ,596 effect of lumbering 596 effect of reforesting 598 function of lnM« .388. 673 grouse as A game crop 587 grouse nianaE''nienl on private hmd 392. .596. 685 grouse managi-menl on public land 596. 682 harmful practices 599 ideal coverts (sec coverts) improving and maintaining grouse habitat 233, 596 INDEX 907 integration with other land use 588, 598. 615. 623, 650 intensive vs. inridental 587 maintenance uf a grouse crap 596. 667 management plans 632 of coverls for hunting 594. 629 organizing the forces of production 629 regulation of hunting 602. 678 regulation of the grouse harvest 631. 673 relation to soil conservation 587, 597 role of the State (see State) significance and potentialities 58], 586 sludieB of 30. 582 summary of pertinent research 583 mandrake 195 manganese, a food component 239, 111 Manitau Island (Wis. J 19 Manitoha 23. 453, 504. 558 Manlius School (N.YJ 30 ManruBs, F. N. 22. 28, 30 man's aids, effect on grouse abundance 233. 370, 392 automobiles and highways IS, 390, 392, 394 axe and plow 387, 393 fire 393 grouse dugs 395 guns, traps and snares 15. 394 methods of taking grouse 8, 9 maple 115. 198. 204, 215. 218, 219, 222. 224. 225, 233 maple, moose (see maple, striped) maple, mountain 204. 207, 215, 220, 225. 227. 655 maple, red 204. 215. 227, 640. 655. 848, 883 maple, striped 204. 215. 225. 227. 655. 848 maple, sugar 190, 204. 228, 2.32, frlO, 655. 818. 8^3 maple family, as a source of grouse food 204 maple-tree worm 101, 212 maps, used by Investigation 695 Marble. 0. B. 78 Maritime Provinces 562 market hunting 14. 388. 394 njcthods 8. 371. 375. 389 numbers taken 8. 371. 375, 558 outlawed 17. 388, 389 market price 8, 101. 371. 557 marking grouse (sec methods and lechniqucd) Murmota nionax 333 Marshall W. 22. 24 marten 333.578 Maries amt-ricana 333, 578 Maries pennanli 333 Marthas Vineyard (Mass.) 18. 50. 507 Martin, A. M. 3 Marvin. C. F. 579 Maryland 2. 13. 18. 23, 24, 36, 46. 72. 272. 274. 420 Maryland Conservation Department 23. 24 Massachusetts 10. 12. 17. 22, 30, 102. 187. 261. 375. 409. 423 421 428. 562, 563, 565. 566, 572 Massachusetts Agricultural College 30 Massachusetts Department of Conservation xxxvi. 22. 24. 30 Massachusetts Fish and Game Division 18. 30 Massachusetts Fish and (;ame Proleclivc Association xxxvi, 30 mast 202 Matanu'k conference 575 Muiheson, R. xxxvi Mathews. F. S. 235 mating habits 67, 248. 265. 282 May, J. B,343 may-apple (sec mandrake) mayflowcr. (Canada 845 Maynard. C. J. 10, 18, 102. 260. 276 McAtee, W. L. xxxvi, 101, 333, 337, 343, .314, 513 McCarthy. D. F. 23 McCormack. J. T. xxxv McDonald, D. 572 McLean. C. P. 22. 24 McNamara, L. C. 23 McVicar. A. J. 22 mealworm 189 Mearns, E. A. 360 measurements 98 mechanical injury as a cause of mortality 402. 511, 528. 531. 532 relation to disease 404 Mecoplera 775 Mi'gninia 409 Melamphy. G. E. 22 melanism 58 Mflanoplus jemur-ruUrum 213 Mennpun 409. 435 Mfphitis mephitis 323, 333- Merrill, A. 22. 24, 26. 30. 447 Mershon, W. B. 10. 32, 520 mesoptiles 59, 746 metabolism 749 methods and techniques used by Investigation 693 analysing chemical composition of grouse foods 237 biometrical analysis of data (see also biumetrical analysis) 718 determining buffer abundance 709 determining disease relationships 711 determining effect of hunting 713 determining food relationships 184, 706 determining predator abundance and activity 708 determining predator food habits 709 determining predators responsible for grouse kills and nest destruction 709 determining sex and age 514, 718 determining shelter relationships 701 determining weather relationships 707 estimating grouse populations (census) 714 making and recording field observations 699. 700 mapping study areas 695 organization 693 principle of random distribution 701 recording observations 699, 702. 703. 704. "05 securing observations 699 selection of personnel 699 selection of study areas 694 trapping and marking grouse 507, 716 methiids of altering existing cover 639 cutting 234, 6-tO fire 228, 232, 612, 650, 664 girdling 640 grazing 233, 641 indirect 59b poisoning 640 methods of artificial propagation (nee aUo artificial propagalitm) all-purpose pens 449. 453. 459, 467. 469. 486 brooder operation 186 brooders and brooder houurs 449. 183 brooding of chicks 419. 182 care of breeding slock 159 care of chicks at hatching time 181 care of chicks following hatching 482 care rtf eggs during incubation 47.) care of young grouse during rearing perioil 491 control of feather picking and cannibalism 188. 190. 497. 501 diseaHc control 435. 437. 438. 139, 4U. 416. U7. 157. 472. 190. 491 egg collection 451. 171 feeding adult grouse 462. 468. 189 feeding chickii and young grouse 187, 4.'>6. 188. 193 handling wild-trapped breeders 419, 452 incubation of grou!>e eegs 449, 473 incubator-brooder method 157 incubators and their operation 475 natural cover meihod 150. 153 obtaining fertile eggs 170 overwintering breeders 159 pens and penning 119. 153. 4,55, 4.59. 461, 492 protection from preilators 4.55, 190 provision of ^heller 4.56 rearing field 191 selection of breeders U9. 163. 196 "eleclivr breeding M»3 sex ratios 170. 878 transportation of eggs 451 use of eggs from wild nesls 151 use of proiise or hens for incubation 1-17. 171 unc of wild-trapped breeders 149 water ref|uiremen1s 162, 482. 489. 491. 501 weight BH an index nf health 463. 500 methods of management altering exi-iling cover (see methods of altering existing cover) artificial food and shelter 632 clear-cut units (see slashings) control of buffers 670 control of deer 666 control of disease 631. 667, 668. 671 etmlrol of livestock .381. 641. 66.=; control of man (see also himting) 629. 631, 678 control of predators .390. 587. 6.30. 667. 668 cover arrangement 609 cover design 607 cover survey and analysis 62 1 cover type mapping 62h determining grouse populations (census) 667, 67,";. 714 forest stand and game cover improvement 623. 650. 777 function of each cover Ivpe 607 harvest of population surpluses (see population surplus) measuring the harvesiable crop 667. 674, 676 planning cover improvenienls 621 planting (see planting) producing and maintaining productive cover (see habitat management) protecting existing habitat 661 restocking (see restocking 1 role of fire 232. 393. 642. 650. 664 role of refuges 391. 629, 667. 679 ' setting up managemcnl plans 632 treatment of open land 6l6 treatment of overgrown land 618 treatuieni of slashings 621 908 INDEX ircaliiK-nl uf woodland 620 U9C of rhrmiral poisons 610. 651 metliutts of taking gruii^tf (see man's aids) mctliylcylisinc 195 Michigan 8. 10. 19. 23. 30. 33. S3. 51, 197. 208. 373. 371. 376. W9. 419. 420. 423. 424. 433. 434. 520. 557. 559. 560. 561, 563. 564, 566. 567. 568. 572, 577. 845. 846. 817. 848. 849 Michigan Conservation Dvpartment xxxvi, 23. 30. 33, 558, 559. 564, 849 Microfilaria (sec also blood parasitrsl 409. 110. 430 Middletown (N.Y.) 203 migration 11. U. 248. 254. 671 Millais. J. (;. 251 MilltT. J. I". 30 iMJnrral muttrr. a food I'lmpoiKMit 236. 237. 239. 244. 815 Mingun Islands (Que.) 50 mink 15. 330, 333. 339, 341, 578, 683, 711, 875 Minnesota 30. 53, 244. 312. 374, 377, 404. 409, 416, 420, 422, 423, 424. 434, 4.36, 520. 521. 524. 559, 560, 561. .'^63. 566. 567. 572. 577, 675 Minnesota Conservation Department xxxvi MiK9issipi)i 52. 53 Missouri 19. 50. 508 Mifisniiri Conservation Cummis<>ion 19 Mitcht'llfi repens 198. 208. 819 mite 409. 410. 417. 4.30 mite, air-sac (foe air-sar niitloBy (Colif.) 47 muskral 341. 417. S78. 683 Mustrla cicognanii 323. .3.3.3. 711 Musteltt noveboracemis 323. 332. 711 Musteta vison 330. 333. 576 Mycobactrrium avium (»ee also tuberculosis) 4.39 Mytmiea 213 Nflgcl. W. 0. 50 nameti of grouse, common 2, 6, 274 names of grouse, scienlitic 46, 17 nannybcrty 653. 662, 849. 8^3 Nantucket Island (Mass.) .SO natal down 79. 748 Naichei (MiM.) S2 Nebraska 50 necrotic stomalitis.*disease of elk 671 NeUon, A. I.. 31. 183, 220 Nelson. E. W. 52. 197 N'clsoD River (Man.) 52. .56 Nematoda 409 ncossoptile 59. 79. 746 nest location (see also nesting cover I 218. 28.i effect of openings 132. 134. 790. 79J effect of slashings 131. 791 influence of conifers 132. 789 relation to drumming In^ 266 relation to exposure 130. 286. 787 relation to slope 130. 786 sites preferred 130. 285. 785 nest mortality (see also survival during nest period) 311 chief predators involved 307. 308. 315, 334, 340. 315. 347. .385. 669 etfecl of predator control 308. 327. 346. 667 number of nests observed 290. 311, 527. 876 proportion attributed to various causes 527 quality of evidence ascribed to diagnosis of broken-up nests ex- amined 876 recorded by other observers 312 recorded during Investigation 135. 307. 311. S25 relation to variations in buffer abundance 307. 315 relation to variations in predator jiressure 314 relation to weather 301 role of predation 307, 311, 526 nest structure 285 nest survival (see also survival during nest period I effect of crown cover 135. 791 effect of distance from opening 136. 798 effect of thickets vs. open cover 136. 797 effect of undergrowth density 135. 796 nesting (see also nest location) number of nests observecl 127. 290 occurrence of desertion 287. 290. 527 occurrence of renesting 248. 291, 364 parasitism by pheasant 273, 290 proportion of females nesting 359, 517 season 248, 281 tolerance of other nesting grouse 290 nesting cover (see also nest location) basic pattern .589 composition 589 extent 589 importance of undergrowth density 128, 589. 781 influence of slope 589 types chosen 127, 783 net productivity (see productivity) Ncuroptera 775 Nevada 50, 52, 54. 196 New Brunswick 563. 566. 573 Newcome. A. 22, 23 New England 11, 30, 53, 54, 183. 220. 221, 232. 340. 343, 396, 408. 409. 415. 416. 419. 420. 433. 560. 562. S6S. 567. 568. 572 New England Grouse Investigation Committee 33, 562 Newfoundland 50. 2.S4 New Hampshire U. 13, 18. 22. 29. 30. 102. 1^3. 293. 401. 109. 422. 434, 557. 561. 563. 565, .S66 New Hampshire Fish and Came Commission .30 New Hampshire F'orcst Service 18, 30 New Jersey 3. 23. 260, 415. 420. 423 New Jersey Board o( Fish and Came Commissioners 23 New Mexico 309 New York 7. 11. 13. 18. 22. 29. 30. 53. SI. 56. 21S. 217. 220. 221. 222, 225, 226. 228. 229. 231, 2.32. 234. 235, 236, 2.39, 241. 242, 244. 245. 265. 268. 273. 278. 285, 310. 312. .323. 324. 325, 326. 331. 333. 337. 338. 340. 369. 372. 373. 374. 37S. 376. 377. 379. 381. 382. 384. .385. 388, 389, 390. .391, 404, 409. 415. 416. 417. 419. 420. 423. 424. 426, 427. 428. 430, 432. 433, 434. 513. 520. 524. 525. 557. 560. 561, .562. 563. S66. 567. 568. 572. 579. 589. 676, 684. 815. 816. 817. 818. 819 New York, province of 17 New York Cilv 8. 10, 17, .56. 273 New York County (N,Y.) 56 New York Forest, Fish and Game Comniifis>iou 12, 13. 30 New York Slate College of Agriculture 31. 182, 598. 687, 688. 749 New York State Conservation Department I6. 18, 19, 22, 24, 30, 381. 417. 563. 601. 683. 687. 688, 721. 719 New York Sidle Constitution 681 New York Stale Department of Public )l<-allli 117 New York Slate Legislature 388 New York Stale Veterinary College 31. UH NickerMin. F. 23 nicotine sulphate 196 nightshade, bittersweet 191. 195. 210. 819 nitrogen-free extract, a fooil component 237. 238. 210, 815 Nitlany Valley (P«.) 845. 816, 847. 819 Noble 18 non. breeding 3.S.3. 355. 525. Ul Norris. L. C. 185. 237. 2.38. 239. 182 Norlh America 38. 426. 430. 432. 5S6. 578 North Carolina 23. 24. 333 Northrillc (N.Y.) 371 INDEX 909 Norwich (N.Y.) 391 Nova Scotia 3, 18, 33. 47, 454, 504, 507, 566. 567 number of eggs, hand-reared grouse 454 number of eggs, wild grouse 71, 354, 360 in first nests 354, 361 in renesls 291, 354, 364. 526 regional variations in New York 362 yearly variations 362 number of grouse in New York, estimatnl 101 number of grouse killed by hunting crippling loss 376 proportion taken 373, 378 reported kill in New York 372 seasonal and daily bags 10, 369, 375 success ratio 369. 372, 375. 376, 378 nutrition (see food of grouse; physiological srinlies) nutritional deficiency relation to brood mortality 528 relation to disease 404. 405. 415 nutritive requirements 238, 239 nutritive value of food, as a contributory influciire in flucluiilinn-* in grouse abundance 241, 575 Nuttall. T. 9, 11. 13. 14. 260. 276. .i57 Nyctea nyctea 324. 333 Nyssa sylvatica 206 oak 198, 202. 215. 218, 220, 221. 228 oak, black 202. 215 oak. bur 202 oak. pin 202 oak, red 202. 205. 215. 640. 655. 846. 883 oak. scarlet 190. 202 uak, scrub 202. 227. 655, 662. 846. 847. 883 oak. while 188, 202. 227. 655. 846, 883 imk family, as a source of grouse food 201 oat 210. 212 Odonata 775 oestrus 67. 267 Ohio 17. 19, 23. 24. 30. SO. 53. .54. 255. 508 Ohio Conservation Department 19. 20, 23. 24 Ohio River 255 Ohio Wildlife Research Unit 30 Oklahoma 52 Oneida County (N.Y.) 114 Ontario 12. 31. 312. 323. 324, 404, 409. 420. 427. 430. 433. 431. 557. 558, 560. 562. 563. 566. 567. .568, .572. .574. 845. 847 Ontario Plain (N.Y.) 36 openings, cover effect on nest survival 136. 798 influence on nesl location 132, 134, 790, 792 man-made 134, 176 relatinn to food supply 231, 776 significance in grouse habitat 132. 170. 596 open l.inil (see also coverj planting design for 616 use hv adults 152, 156. 819. 821 use by broods 140. 801 use for nesting 128. 783 ..possum 15. 324. 330. 333. 334. 310, 342. 344, 669. 711. 875. 876 Orange County (N.Y.) 287 orchard damage by grouse 102, 187 Ord. C. 47 » Oregon 48 Oregon Wildlife Research Unit 19. 20. 509 Ornithoica vicina 409 Orobancliaceae 210 O'Roke, E. C. 433 Orthoptera 198. 212. 223. 775 Osborne, L. xxxv osprey 333 Oslrya 198 Ostrya virginiana 186. 201. 846 Oswego County (N.Y.) 210 Otsego County (N.Y.) 431 Otselic valley (N.Y.) (iQS Ott. F. 23. 24 OtiiS asin 333 Ouachita Mountains (Ark.) 50, 52 overgrown land (see also cover) design 618 maintenance 641 use bv adults 1.52. 155. 819. 821 use by broods 140. 801 use for nesting 128. 783 owl (see various species) 14. 309, 334, 335. 337. 391. 432. 439. 870 owl. barred 330. 333. 337, 339, 342. 344. 875 owl, great grav 333 owl, great horned 308. 313. 321, 323. 324. 329. .3.30. 333, 334. 336. 338. 339. 341. 344, 346, 381. 432. 552. 630. 667. 668. 669, 875. 876 as a predator of adult grouse 27. •308. 322. 328. 337. 348. 455. 876 frequency of grouse in pellets in relation to buffer abundance 322 owl. long-eared 875 owl. saw whet 875 owl, screech 57. 333. 342. 875 owl, short-eared 875 owl, snowy 330. 333. 337, 339, 343. 875 periodic influxes from the North 321 oxalic acid 195 Oxalis 195. 210 ' Oxvspirura petrowi 409. 434 Ozark Mountains (.\rk.-Mo.) 4. 50, .393 Palmer. R. 339 Pandion haliaetus 333 panther 309. 333 paralysis, relation of vitamins to 239 parasites, animal (see also various species) 405 manner of infection 408 numbers encountered per bird 411. 414, 416. 418. 421, 42i. 426. 432 regional dislribuliun 401. 409. 412, 413. 415, 416. .574. 575 relation to age of bird 410. 412. 413 relation to disease 403, 404 relative occurrence in New York IJO. 115, 430 seasonal variations 412. 413 species found in grouse 409 yearly variations 413 p.irasites. plant 404. 408 liarasitium and disease 401, 410. 694 animal parasites of grouse (see also parasites) 405 de of 310. 511. 513. 551 significance of .109. 331, 338 under artificial propagation 4SS predator control 345 effect on grouse productivity 308. 350 effect on predators 346. 552 experiments ctmducled 315 number of predators taken 329. 315. 875 place in a management pr..gr(im 307. 309. 345. 350. 630. 667. 6(.8 results during adult period .308. 317. 349 results during brood perioil 308. 347, 349 results during nest period 307. 308. .327. 345. .316. 349 selective 331. 345. .3.50. .5.52. 667. 669 ]i4 INDEX 911 numbers taken during control experiments 329. 8T5 principal species in New York 329. 338. 667 relative importance of various species during adult period 308. 337, 338. 669 relative importance of various sjiecics during Iirood period 308. 335, 338 relative importance of various species diiriny nesi period 307. 334. 338. 669 species of the Northeast (see also various species) 308. 332 variations in distribution in New York 324, 330. 337. 344 predators, food habits of (see also various species) 338 methods of study 338. 709 seasonal distribution of stomachs examined 875 presidential elections 579 prices, of grouse (see niarkel price) primary fealliers 60. 743 Prince Edward Island 50, 251 private lands, relation to grouse managenicnt 685 Procyon lotor 323, 333 productivity, net (see also life equations) 332, 511. 512. .5.i9. j17 effect of controlled population reduction 522. 538, 541 occurrence of surpluses 539. 550 production ratios (April) 542, 543 production ratios (September) 511. 539 reasons for low productivity observed 539. 544. 572 relation to density of breeding population 350, 511. 539. 540. 541. 542 relation to range quality 512. .522. 5.35. 516. 551. 553. 668 relative significance of losses during different life periods 332. .539. 510. 541. 542. .'^43. .544. 51(.. 547. 518. 519. .571 trends observed by Invesiif-ation 512. .523. 539. 513. 544, 54S. 882 productivity of grouse iM.pulati.ms («e.- al«o reprodurlivc capacity ) 511.525 breeding success 353. 513, 517. 525 definition of term 525 interrelationships involved 511. 512. 513. 528. 532. 535. 541. 516. 550, 551, 552. 553 life equations (see life equations) net proiluclivily (see proiluclivity. net) iqiportunity l"r management 512, 521. 546, 553. 577 jiotential 309. 512. 545 surplus (see population Burplut) survival (sec survival during adult, brood and nest periods) promiscuity 248. 266. 267. 3.55. 368 propagation (see artificial propagation) Proslhogonimus macrorchis 409, 434 Protection Island (Ore.) 19. 20. 509 protein a food component 236. 237. 845 proportion needed by young grouse 185 protozoa 409 proventricular roundworm (see sloniarh wotm) Prunui 186. 198. 199 Prunus pennsyltanicum 847 Prunus scrotina 847 Prunus virginuma 847. 848 Pseudolynchia canaiiemis 433 PseudoHcorpionida 775 psycliology 63, 247 ptarmigan (see also grouse, red; grouse, willow I 16. 427, 128 j.terylography 59. 78, 741 age variations (see also plumage develoinunit ) 7 IK brood spot 746 feather spaces 59. 711. 745 feather tracts 59. 741 numbei of feaihers 36. 60. 717 plumage wear 747 "snowshoes" 741, 748 types of feathers 59, 746 PtYchostomii bonasae 409. 433 public hunting grounds 391, 684 public lands, integrating giouse management on 682 Pugct Sound (Wash.) 18 pullorum disease 415 pulmonary mycosis (sec aspergillosis) Pnpiparia 432 pussey's toes 198. 204 Putnam County (N.Y.) 18 Pyrola 203 quail 46. 73, 74, 229, 235, 267, 327, 415. 416, 419. 420. 435. 436. 437. 438. 674 quail, bobwhile 46. 267. 327. 394, 416, 418. 423. 426. 428, 434. 849 rjuail, valley 433 ■■([uail disease" 402. 416, 435. 436 cause 436 control 437. 450. 472. 492. 495 dissemination 437 nature 436 Quebec 17. 33. 47. 56, 409. 128. 557. .566. 567 (Jueen (iliarlottc Islands (Br. Col.) .52 Queens County (N.Y.) 56 Quercus 198. 202 Quercus alba 202. 846 Quercus borealis 202, 846 Quercus coccina 202 Quercus ilicifolia 202. 846. 817 Quercus macrocarpa 202 Quercus palustris 202 Quercus stellala 849 Quercus veluiina 202 rabbit (sec also buffers) 14. 308. 309. 322. 329, 339. 340. 311. 342. 343. 344. 552, 709 rabbit, cottontail 192, 322. 325. .339. 340. 345, 430. 670. 682 as a source of tularemia infection 416, 417 fluctuations in abundance 326. 327 rabbit, jack 430 rabbit, snowshoe (see varying haret raccoon 15. 313. 323. 329. 330. 333. 334. 336. 337. 338. 339. 340. 341. 344. 382. 456. 496. 669, 683. 711. 875. 876 Radi. M. H. 78 Rae, T. 22. 24 ragweed 206. 8^19 Raillietina tetragona (see also large tapeworm) 409, 412. 426 rain, as a source of drinking water 243 rainfall (see precipitation ; weather) rainwaler. pre-nence of mineral matter in 214 Randall. P. E. 314 range (see also di^^tribulion) 36. 48. 49 doubtful records 52 on various islands 50, 254 present 18, 49 primitive 18. 49 shrinkage of 50. 51 sporadic occurrence 52. 56 Kanunculaceae 208 Ranunculus 208 Ranunculus ocris 847 Kaxmussen. D. i. xxxvi. 52 raspberrv 182. 189, 198. 199. 207, 214. 215. 216. 218. 219, 220. 221. 222. 223. 224. 225. 227. 243. 707 raspberry, black 847 raspberrv. red 817 ral. .Norway IS. 311. 343. 456. 491, 192. 496 rate of spreatl of grouse populalions 512. S22 raven 333 Rawson. C. L. 22. 24 razorbsck (sec hog I Red River (N.Y.) 204 reflunbes flintoncr from previous flunh 167. 813 relation to cover type 166. 842 rcforeotation areas 174, 598. 612, 616, 683 refuge, national waterfowl refuges (sec also methods of management) 388, 391 establishment 20 relation to grttuse abundance 391 seed stock 603. 629 value of 20, 667. 679 regions of New York, description 113, 596. 694 Adirondack 114. 596, 694 Calskill 115, 596. 694 rest of State 116, 597, 694 regression (see biometricat analysis) Reiner. N. E. 15 relative abundance of grouse 53. 55. 676 rencsting 218. 291. 354. 364. 519, 525 reported kill of grouse in New York 372 reproductive capacity (see also eggs; embryology; number of eggs) 30, 353. 551 effect of age 359 fertility of eggs 354. 365. 454. 526, 527 inbreeding 354. 367 bmgevily and senility 360 n on -breeding 355 number of eggs laid 354. 360. 560 renesting 354. 364. 519. 525 sex ratio 353, 355 viability (hatchabilitv \ of eggs 72. 354. 365, 527 reproductive potential 309. 354. 511. 512. 514. 545, 553, 571 requirements of gralmon, ,\ll.inlic .'>78 :iamburus 204 Sambucus cunudensis 204 Sambucus riHcmosa 201 Samnels, E.A. 286 sanctuaries 391 Sandys, E. 235 Saratoga (N.Y.) .371, 557 SaratoRfl Hotel 12, 557 Saskalrbcwan 18. 48, 50. S\. 258 sassafras 212 Sassafras vanifnlium 212 ftnluration point 330. 512, 521. 668 SaunilctB. t;. It. 373, 376 saw-fly lOl. 212, 213. 223. 224, 232 law-fly. larch 232 Sawyer. K.J. 274. 278 Saxifragn 20(> Saxifragaceac 206 saxifrage 206 naxifragr (anuly. an a nnurco u( gruuse food 206 Scandanavia S78 Hinrcity (sec also fluctuatlona) 693 . 816. 817. 8IH. 819 early records xxxi. 11. .5.S8. .560. 562. 563 remedies that have been proposed 17. 20 surveys and suggested causes of 12 Schoharie County (N.Y.) 204 Srboolcraft County (Mich.) 845 Schuyler County (N.Y.) 695 Sciurus carolinensis 326 Sciurus kudsonicus 327. 333 Scluter. W. I.. .SO Scoliaptex neiulosa 333 Scotland 103. 2S». 630. 671 Scott. T. G. 339 Scudder, B. A. 15 Seamans, R. 31 scas^onal activities 250 si-asnns. open (see bunting regulations) second broods. lack of 248. 291 second nesis (sec rcnesting) secondary feathers 60. 713 sedge 188. 198. 203. 207, 218, 219. 221. 222. 223. 224. 22.'^. 226. 227. 231 sedge, bladder 203. 215, 222. 815 sedge, bristle-stalked 215 sedge, graceful 215, 222 sedge, green 222 sedge, pale 215 sedge, sallow 845 sedge, sickle 215 sedge, slender-stalked 203. 215, 222 sedge family, a!^ a ouurce of grouse food 203 seeds, used in place of grit 192 selective cutting frl4. 651 semiplumes 59 senility 360 service bcrr> (see shadbush) Setaria lutescens 845 Scton, E. T. 256, 325, 326. 333. 536 sex influence on flight behavior 105, 167. 843 recognition of 39, 167. 514. 718 sex ratio 15. 267, 514 among newly hatched chicks 353, 355, 514 of adult specimens collected during winter 515 of immature specimens collected 516 seasonal trend 353, 355. 512, 516 summer relationsbiji on study area 355. 514 sex rhythm 68. 267 shadbush 190. 198. 200. 205. 209. 210. 211. 2iS. 218. 219, 220. 222, 224, 225. 227. 640., 655, ai7. 883 shelter (see also cover; tables 121 to 171 in Appendix) 30, 105. 511, 513 relation to carrying capacity 523. 551 shelter relationships, determining 701 shelter value of cover, recording 698 Sherburne State Game Farm (N.Y.) 28 Sherman, I., ft. 22 Shillinger. J. E. 415. 116. H7. 133. 136. 568, 574 shinleaf 203 Shipley. A. E. 427 shooting, spring and summer 17 shrew (see also buffers) 308, 322. 325. 327. 329. 333. 339. 340. 341. 342. 313. 670. 709 shrubs (see trees and shtiibs) Shufeldt, R. W. K.. 725 Sbutt, F. T. 845.847. 819 Sibley. C. L. 24 Simulium venustum 433 Sines. H.J. 22 Sitka (Atsk.) 52 size 37. 98 skeletal measuremcnta 727 skeletal structure 721. 723 Skill. J. \ . XXXV, 262, 395 Skinner. M. 1". 333 skull (see anatomy) skunk 11, 32,1. 324. 329. 330. 333. .334, 337. 338. 339, 310. 31 342, 314. 381. 382. 552, 669. 683, 711, 875, 876 skunk cabbage 210, 815 slashings (see also cover; openings) design of 621 influence on nest location 134, 791 place in a nianagemrnt program 176, 617 suggestions for establishments of 647 use by adults 152. 1.55. 819. 821 u«o by broods 140. 801 UBO (or neslinR 128. 131. 783 value of clear-cut areas 176. 233. 647, 776 flipped tendon (see pcrosis) •lope relation to nest location 130. 786 use by adultit in relation to time of day 16'!. 6^f8 use by adults in relation to weather Ifrl. 839 use by broodtt in lelaiion In ogc 148, 813 INDEX 913 use by broods in relation to time of day 149, 814 use by broods in relation to weather 149, 815 slug 408 smartweed 194. 222, 847 Smilacina racemosa 845 Smilax 210 Smilax rotundifolia 845 Smith, R. G. 23 Smith. T. B. 292 Smith. T. J. 438 Smith Landing (N.W. Terr.) 285 SmithviUe Flats (N.Y.) 558 Smyth. T. 30. 33. 98. 183. 194. 201. 203. 255. 269. 285, 339 snail 434, 671 as a source of gapeworm infection 418 as a source of grouse fond 192, 198, 213. 221. 223. 224 as a source of tapeworm infection 408 snail, pond 214 snake 15 snake, garter 192, 342, 344 snake, green 188, 192 Snake River Plains (Idaho) 48 snares (see man's aids) snaring grouse 14, 252 snow (see also precipitation; wt-allifrj as a source of drinking water 2U snow roosting 163, 248. 302 snowberry 204 "snowshocs" 48. 63, 248. 269. 270 Snyder, L. L. xxxvi social order 37. 63, 268 among hand-roared grouse 416. 162, 191. 196 relation to rex 64 significance in wild 65 sodium arsenitc, use in poisoning trees and shrulm 6t0 soil eflect on mineral content of plants 239 relation to rover and habitat 169 soil conservation district 598, 686. 687. 688 solanine 195 Solarium Dulcamara 194. 210. 849 solar activity (sec ulsn cyrlcB) as a contributory influence in fluituuiinns in grouse abundance 575 relation to other natural phenomena 575 relation to weather factors affecting food 242, 575 solarization 242 Solidago 204 Solomon's seal 845 Solomon's seal, false 845 Solomon's seal, two-leaved 220 Soper, J. D. xxxvi sorrel 235 sorrel, sheep 195, 198, 206, 220 sorrel, wood 195, 210, 218 sour gum 206 South America 426 South Carolina .'".2. 197. 210 South Corver (Mass.) 26 South Dakota 50, 333 Southern New York Fish and Game Association 18 Southern Tier section (N.Y.) 56, 104, 116, 691 sowbug 414 aa a source of stonuirh worm infection 407. 408. 415. 420, 491 sjiarrow 418, 439 sparrow, house 420 speedwell 190 Sperry. C. C. 33 Sphaerium 192 spiders 198. 212, 213, 221. 223. 221. 110. 776 spike-rush 192 Spiller. li. L. 2.30. 25,5, 2(iO Spoare. J. W. 12 Sporting Arms and Ammunition Manufuctuicr's Institute 30 sporting value of grouse 20. 103, 371 sprays, used by oichardtsls and foresters 196 spring nesting grounds 127, 153, 160, 64-t, 783, 825 spring shooting 10 spring shuffle 248. 256. 348 spruce 115, 210. 235, 386 spruce, black 56 spruce, Norway 655, 883 spruce, red 655. 883 spruce, white 655. 883 squirrel (see also various species) 15. 325, 327. 343, 709 fluctuations in abundance 326 squirrel, flying 341 squirrel, gray 57, 326, 340, 341, 683 squirrel, red 327. 333. 334. 340, 341, 381. 391, 552. 670. 710. 87, 876 Slaked Plains (Tex.) 52 Staples, E. F. 10, 14. 371 starch, a food component 238 starvation, effect uf (see physiological studies! State, role of in grouse management 601 acquisition and development of grouse lands 603 annual inventory 602 maintenance of hunting grounds 604 regulation of hunting 602 research 602 service to landowners 603 Staten Island (N.Y.) IB statistical analysis of data (see biometrical analysis) Stephens. J. F. 46 sternum (see anatomy) Stpwart. R. 31 stink-bug 195. 213. 224 stock, common 578 Stoddard. H. L. 267. 509 Sioddart. A. M. xxxi. 15. 17. 30. 367 stomach inflammation 411 stomach worm 404. 406, 419. 435, 571 description 420. 425 distribution 402. 409. 412. 413. 415. 416. 420 incidence and importance 402, 410. 411, 412, 413, 415. 416, 417, 422 life cycle and dissemination 407. 108. 414. 420. 494 pathogenicity 407. 411, 421 stratagems of alarmed grouse 262 strawberry 186. 189. 190. 198. 199. 205. 207. 209. 215, 216. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 231. 233. 707 strawberry, barren 198. 200. 211. 215. 218. 220 strawberry, wild 847 strawberry worm 409, 434 Strix varia 330. 333 utrnngilosis 630 strutting 282 as a part of rourlship 65. 282 relation to mainienanrr of Mtrial itrder 64. 282 strychnine 196 study areas used bv Invef^ligntion. drdcriplinnp Adirondack 695. 696 Catskill 695 Connecticut Hill 695 606. 6'»7 Pharsalia 695 Slurgis 18 subclimax forest tsvr f.ii.-ii) subspecies characters 48 classification 47 discovery 47 distribution 47. 49. 53 Suhulura 409 succession, forest after cfear-cuiting 177 effect of fire on 612 * relation to grouse food plants 231 role of lis succulence, 09 a source of w.iler 243 sugar, a food component 238 Sullivan County (N.Y.) 213 sumach 196. 198. 199, 215. 216. 217. 218, 219. 220. 225. 226, 238. 239. 243. 707 sumach, dwarf 215. 227. 655. 662. 883 Humach. smooth 211. 215. 227. 655. 662, 848 sumach, slaghom 186. 190. 203, 205, 209, 215, 227. 2.^7. 239. 243. 244. 655. 662, 848. 883 sumach family, aa a source of grouse food 203 summer boarders 15 summer feeding grounds 1.19. 1.52. 1.55, 2.30. 617, 801. 819. 827 sunflower 849 sunlight, as a weather factor affecting food composition 242 sunspots (sec solar activity) Superior National Forest (Minn.) 371. 521 surplus (see population surplus) surveys (sec also methods and techniques) of others regarding grouse 12. 30, .32 survival during ndult period (see also aN of murlalily 307, 308, 526, 527 Su»quchanna River 57. 254 Suttim (Mu»8.) 25. 26 Siiilrm. G. M. 15. 2,W, 333 SwainHon, W. 6 Swreli;rn»!i HilU (Muni.) 50 HwiiiimiiiK, by grouse 252 Switzorlun.) 431 Sylvilagu.s flondanus 322 Symphorirarpos 204 Syniplocarpus foetidus 210,8-45 Stngiimus 408 S\ngamus Iracht-n (src also {;ii|>i"*»orrn ) 409, 416. 417, 418. 425 »yiisa»Tuni («<■«• anatomy) Syracuse (N.Y.) 12 Syrarusr I iiiv«T»ily 30 syrinx (see anatomy) Syfllema Naturae 46 Taintor. C. M. 18 tamarack 56. 845 tatiK-nefis 260, 262 Tamias striatus 333 lannin. a fuitd romjMinent 202. 238 ta|.<-w..riii 107. 108. KW. 11(1. 112. Mr,. 119. 127. 4.34 lajicwuriii. lai^f itrHcription 426 distribution 413, 426 incidence ami inijiorlunce 410, 412. 413. 427 life cyrle and dissemination 426 pathogenieily 412, 427 tapeworm, small descri|>tion 427 distribution 413. 427 incidentre and iniportanrr 410. 412. 413. 427 life cycle and disseminati<»n 427 Iialhiigenicity 412. 427 tar, a fi>od component 239 Taraxacum officinale 849 Taunlon (Mass.) 371 taxine 195 taxonomy 38 Taxus canadensis 194 Tuylor, E. L, 418 Teal. H, A. XXXV teleoptilcs 59, 746 temperature, air (see also pbysiologiral studies: weather) relatioh'to f.dnlt mortality 304 relation to breeding in sheep 576 relation to brood mortality 299. .303, 576 relation to cover chosen (see also cover) 142. 147, l6l. 305, 804, 831 'relation to nesting dales 299, 303 relation to plant mmposilion 242 relation to population fluctuations 299, 305 trniperaiuH-. iM.iiy I sei- also physiidocical studies) 37 Temporary Emergency Relief Administration xxxvi Tennesse.-' 36. 46. 52. 272. 123 Trntliredinidoe 213 terrestrial magnelism 578 territory (see alnu liahiiat) 248, 257 a|>plieation of term 257 of adults in hrerdinc season 257. 368 of broods 294 relation to eover quality 257. 259 seasonal shifts 259 size 258, 273 year around 257, 506 Tftramem americana 409, 431 Telramorium 426 Tetrao 46 Tvtrau fusca 47 Teltao snitini 47 Tftrao fogaius 47 Trtrao V mbelloidi'S 47 Trirau Umhftlus 46 Trtraonidae 46. 721. 724, 725. 727 Texas 52. 420 thickets, elfert on ne».t survival 136, 797 Thon-au, H. 1). 21') thornapple 182. 188. 189. 190. 196. 198. 200, 207, 209, 210. 2U, 21.5, 217. 218. 219. 220. 221. 222. 225. 226, 227. 228. 230, 231. 234. 2.';9. 595, 655. 662. 707, 817. 883 Thornton. M. K. xxxvi, 091 Thotr I. K. U. 657 thruoh 418 Thiirin. (Sn.l.l 254 Thysanoptera 775 TiarrUa cindifulia 206 tick, bird 402, 409. 414, 416, 417. 430. 131 tick, rabbit 409, 4.3U as a sourre of tularemia infeiliitn 416, 117, 430 licks 101. 409. 130 dcsrription UO. 431 distribution 109. 413. 130 ineidrnee and importance 410, 411. 113. 41 1. 432 life cycle and dissfmination 431 pathogenicity 431 Ticondcroga (N.Y.) 3 Tidyman. W. 31, 203 timber production .587, 590. 610. 6.S0. 633. 68:{, 685 time of day r«-Iation to ei.ver chosen 112. 159. 803. 823 relation to slope chosen 149. 164. 814, 838 Todd. W. E. C. 48 tolerance of other species 273 Tompkins County IN.Y.I 199. 212. 373. 376, 378. 433, 520. 695 Tnrrev, H. A. 22, 24. 26, 27. 30. 447. "1.53 Torrry. J. A. 22 touch-me-not 208 Townsend, C. R. 20. 338 Townsend. M. T. 320 Trainer. J. E. 31, .i9, 694, 711 trapping grouse (see methoils am! techniques) trapping predators (see hunting predator;^: predator enntroll traps (see also man's aids) 15. 389. 392. 669 Traiitman. M. II. 23. 24 tree roosting (see roosting habits) trees and shrubs rrop. wolf and weed 623. 777 data regarding propagation for planting purpits*-*, 883 data regarding use» and site requimoriils 6.55 trefoil, tick 208 Trematoda 109 Trichomonas hniui.snr 109, 433 Trichomonas gallinarum 433 Triehoplera 775 Trifolium hyhridum 208. 848 Trifotium pratvnsv 208. 818 Trifolium repvny 208. 818 Trippensee. R. E. 23. 31. 371. 377. 521 Trombicula microti 409. 417, 434 Trypanosoma 430. 433 Trypanosoma giillinarum (see alsn blond piirasitet,) 109 T.uiga ranadt-nsis 210. 8'15 Tubbs. F. F. 557, 562 tuberculosis 404. 408, 416, 436 cause 439 control 439 dissemination 439 nature 439 Tufls. H. W. 451 tularemia 416. 417, 430, 574 tumor 411 turkey 46, lUl, 273. 383. 418. 419, 420. 426. 428. 430, 431, 436. 438 Turrill. D. 22, 24 Tussey Mt. (Pa.) 848 Tnttle, H. E. 737 Tympanuvhus 16. 724, 725, 726, 727. 737 type localities 46. 47 type specimen, of species 3, 46. 47. 274 type specimens, of subspecies 47 types of cover (sec cover) lyzzer, E. E. 33, 429. 438 ulcerative enteritis (see also "quail disease") 435, 436 L'lmus 212 ultra-violet radiation 578 undergrowth importance as brood eovfr 146. 808 importance as nesting cover 128. 784 undergrowth density. ctTect on nest survival 135. 796 imderplanting 620. 661 L . S. llureaii o[ Ainmal Indiislrv xxxvi, 33. 714 II. S. Bureau of Uiologieol Survey 27. 30. 33 V. S. ExperiuK-Mla! Fur Farm 30 U. S. Fish and Wihilife Seivice xxxvi. 31, 1^3. 184, 339. 505. 670. 706. 749 U, S. Forest Service 31, 640 U. S. Geological Survey 626. 696 U. !^. National Museum 72 L, S, I'ublic Health Servi.e 30 IJ. S. Resettleriieut Adniinintralion 520 i:, S. Soil CoHMTvalioM .S|■|^il•e 657, 687, 688 V. S, Weather liur.aii .300. 303. 305. 306, 707 I'niversity of Alberta 520 Uiiivemily of Maine 31 t'niversily of Minnesola 30 I'nivcrsily n to productivity 511. 513, 551, 552 relatiim to slope use 164 Weed. C. M. 203. 206 weevil 212. 213. 224 weevil, black vine 101. 212 weevil, clover root 101 weevil, strawberry 101. 212. 213 Wehr. E. E. xxxvi. 418. 419. 424 weight (see also physiological studies) 37. 90 age differences 91. 94 critical zone 37. 95. 96 danger zone 95. 96 extremes 95 factors affecting 37, 90 loss during egg laying 361 normal variation 91 rif band-raised grouse 463 regional differences 94 relation to health 37. 95 seasonal trend 91 sex differences 91. 92 significance 90 yearly differences 91 Westchester Countv (N.Y.) WcMfield (Mass.) ' 10 West Virginia 23 W.im..re. A. 1. 16. 18 Weimore. P. W. 436 wheat 210. 212 While. J. A. XXXV While Mountains (N.H.) ,54 White Pine Mountains (Nev.) Whitehead. F. E. 8. 196 whortleherrv 2.35 Wight, H. M. 266 willow 195, 201. 218 ^willow, pussy 201 willow family, as a source of grouse fi Wilgu». H. S. 141 Wilson. A. 6. 7. 194. 2.35. 259. 260. 292. Wilson. K. 721 wind (see weather) Wing. I.. W. 5,56. 562. 575 wing (oer anatomy I winter shelter 1.53. 1.59. 616. 823 winierbrrry 210 winterpreen 203. 218. 220. 225, 2.35 w ire-worm 213 Wisconsin 18. 20. .33. 53. I&3, 197. 208 120. 423. 424. 524. 557. 560. 563, 566, 567. 568. 571, 575 Vt isconsin Conservation Dcparlment xxxvi. 5,57" witch-hazel 190. 208. 222. 847 Miichhopple 231 withe-rod 203, 201. 2I.'i. 227, 849 w.ilf 308. .3.33. 578 W ....d. F. W. 22. 21 Hoodchuck 333. .334. 310. 669, 670. 711. 875. 876 Moo.Jc.Kk 394. 6^3 Woodhaven (N.Y.) .5.58 woodlands (see also cover) design 620 use by adults 152. 819, 821 use by broods 140. 801 use for nesting 128. 783 w(Mid. using industries, importance of 598 Woodruff. E. S. xxxi. 12, 13, 14. 15. 30. .568. 572. 571 Woodward. G. K. 719 Worcester (Mass.) 21. 375 Works Progress Administration xxxvi wren. Carolina 420 Wright. T. 845. 817. 818, 849 Wyoming 50, 53, 671 Yale School of Fore.*trv 31 Yapp. R. H. 512 Yeatter, R. E. 50 yew 191. 195, 222. 224 yolk sac 74, 76 York Factory (Man.) 52. 56 Yukon Terriiorv 47. 54. 235 Yukon valley 17. 48. 56 18. .532. I. 542. »d 201 371. 389. 396 212. 220. 221. 326, .375, 118. 161. 305, 801. 810. Zea mays 845