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Natural salt licks as a part of the ecology of the mountain goat Hebert, Daryll Marvin 1967

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NATURAL SALT LICKS AS A PART OF THE ECOLOGY OF THE MOUNTAIN GOAT by DARYLL MARVIN HEBERT B.Sc, University of B r i t i s h Columbia, 19-65 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n the Department of ZOOLOGY We accept th i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA May, I 9 6 7 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced deg ree a t t he U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and S t u d y . I f u r t h e r ag ree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by the Head o f my Depar tment o r by h.i>s r e p r e s e n t a t i v e s . It i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l no t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Depar tment o f The U n i v e r s i t y o f B r i t i s h C o l u m b i a Vancouve r 8, Canada Date i i ABSTRACT The role of natural earth l i c k s i n the ecology of the mountain goat (Oreamnos americanus ( B l a i n v i l l e ) ) was studied during the summers of 1965 and 1966 i n the Rocky Mountain Trench of southern B.C. The patterns of movements of the animals were determined as they used the l i c k s and the vegetation, l i c k s o i l s and blood serum were analyzed with respect to sodium content. The predicated use of l i c k s as suggested by other workers involves the idea that sodium may be the att r a c t i n g element. The present study examined seasonal and d i f f e r e n t i a l patterns of movement, along with p e r i o d i c i t y of use, i n order to determine the resultant interplay of animal movement and sodium and water content of the vegetation. The goat encounters such a d d i t i o n a l r i s k s as predation, parasitism, hunting and joint use while using the l i c k seasonally. Since the goats use the l i c k d i f f e r e n t i a l l y i n time, each sex i s affected to a d i f f e r e n t degree by the above f a c t o r s . D i f f e r e n t i a l use also produces c h a r a c t e r i s t i c patterns of grouping and molting. P e r i o d i c i t y of use occurred mainly i n the afternoon, although distance t r a v e l l e d and complexity of the l i c k may affect time of entry, length of stay and group s i z e . The frequency of use d i f f e r e d at each l i c k , with the average time i i i of use by an i n d i v i d u a l being one to three weeks. Environ-mental factors such as temperature and weather appear to regulate the movement to and from the l i c k . Analysis of the vegetation revealed that sodium was extremely low and that potassium was present i n s u f f i c i e n t quantities to meet the requirements of the animal. No s i g -n i f i c a n t trends were found to exist from spring to f a l l or due to changes i n elevation, as far as sodium and t o t a l ash were concerned. Observations indicated that animals select c e r t a i n l i c k s over others and select s i t e s within a l i c k . These high l i c k i n g s i t e s were shown to have a higher sodium content. Newly established l i c k s had a higher sodium content than did old l i c k s , however, highly preferred s i t e s were not always higher i n calcium, phosphorus or cobalt. A normal range of serum sodium values was> established for the goat but due to the regulatory function of the kidney, changes i n serum sodium due to l i c k use could not be detected. Serum sodium decreased with age. A female with a k i d had a low serum value. It appears that the l e v e l of deficiency causing the craving i s not s u f f i c i e n t to show up i n serum analysis. Animals died during trapping operations and a selenium deficiency was suspected. Gross symptoms approximated those attributed to white muscle disease and the vegetation iv contained selenium i n amounts which are known to cause th i s myopathy. V TABLE OF CONTENTS Page ABSTRACT i i TABLE OF CONTENTS v LIST OF TABLES v i i i LIST OF FIGURES x ACKNOWLEDGEMENTS x i i i INTRODUCTION 1 DESCRIPTION OF THE AREA 5 METHODS AND MATERIALS 9 Trapping 9 Marking 10 Vegetation 11 Lick s o i l s lk Blood 15 RESULTS 16 LICK STRUCTURE 16 T r a i l systems 16 Size and structure 19 PATTERNS OF MOVEMENT AND LICK USE 20 Introduction 20 Seasonal patterns 22 Succulent spring vegetation as a de-terminant of seasonal patterns 27 Seasonal patterns invoke a d d i t i o n a l r i s k s , - 29 I n t e r s p e c i f i c j o i n t use controlled by seasonal patterns 31 INFLUENCE OF SEASONAL PATTERNS ON EXTERNAL 'PARASITE LOAD. PRE DAT I ON AND HUNTER KILL 3^ Predation 3!+ E f f e c t of the hunting season on l i c k use 35 v i Page PATTERNS OF DIFFERENTIAL USE IN TIME 37 D i f f e r e n t i a l use as a general trend 39 D i f f e r e n t i a l use between l i c k s The d i f f e r e n t i a l e f f e c t of parasites, joint use, predators, and hunter k i l l ^1 GROUP SIZE IN THE LICK AS INFLUENCED BY DIFFERENTIAL USE ^3 T o t a l group size kk Male group size *+5 Female group size k6 BEHAVIOUR 50 DIFFERENTIAL USE AND ITS EFFECTS ON MOLT 52 LICK USE OVER A 2*f HOUR PERIOD 56 SEQUENCE OF LICK USE DURING THE SUMMER 58 DISTANCE AS IT RELATES TO LENGTH OF TIME SPENT IN THE LICK 60 Complexity of the l i c k 61 PATTERNS OF FREQUENCY OF USE 63 Duration 6k Frequency and patterns of movement 66 The importance of region 2 67 Environmental factors a f f e c t i n g movement 69 Changeover »in the population and the factors regulating i t 75 Population estimate . 77 VEGETATIVE SOURCES OF MINERALS 78 Introduction 78 Determination of major forage species of the goat 86 Mineral content of the forage 88 LICK SOILS 97 Introduction 97 High and low l i c k i n g s i t e s 98 v i i Page Mineral content 99 New l i c k s 101 SODIUM IN THE ANIMAL 105 Introduction 105 Comparison of sodium values of animals moving into and out of the l i c k 109 SUSPECTED SELENIUM DEFICIENCY 113 DISCUSSION AND CONCLUSIONS 117 LICKS AS A MANAGEMENT TOOL 122 SALTING 122 SUMMARY 125 LITERATURE CITED 130 APPENDIX 136 LIST OF TABLES v i i i Page 1 A monthly comparison of average group s i z e , using data from Brandborg (1955) and data from the East Kootenay region 49 2 The p e r i o d i c i t y of d a i l y use at the Lazy Lake and Dutch Creek l i c k s , based on observations of groups 56 3 The change i n degree of l i c k use, using com-bined group observations from the Dutch Creek and Lazy Lake l i c k s 59 h The average time spent i n the Lazy Lake and Dutch Creek l i c k s based on observations of female groups 6Q> 5 Marked animals and subsequent sightings at the Stoddart and Toby Creek l i c k s during the summer of 1966 65 6 The number of animals observed i n regions 2 and 3 during a period when a storm and tem-perature are out of phase 71 7 The movement of animals between regions 2 and 3 at the Toby Creek l i c k as a r e s u l t of a severe storm 72 8 The movement of animals between regions 2 and 3 at the Toby Creek l i c k as a r e s u l t of a moderate storm 73 9 The e f f e c t of a storm on animal use days of regions 2 and 3 73 10 The comparative use by big game animals of mixtures offered i n sixteen mineral caf e t e r i a s i n western Montana for a two year period (1951 - 52) from Stockstad (1953) 81 11 The comparative use by big game animals of f i v e s o i l impregnation tests i n western Montana, for a two year period (1951 - 52) from Stockstad (1953) 82 12 Rumen analyses from f i v e mountain goats, showing the percentage of forage types 87 ix Page 13 Forage samples co l l e c t e d i n September, 1966, i n regions 1, 2 and 3 at the Toby Creek l i c k , showing the amount of ash as a percentage dry weight 90 ih Forage samples co l l e c t e d i n the spring, 1966, i n regions 1, 2 and 3 at the Toby Creek l i c k , showing the amount of ash as a percentage dry-weight 9 1 15 Forage samples c o l l e c t e d i n September, I966, i n regions 1, 2 and 3 at the Toby Creek l i c k , showing the sodium and potassium concentrations as a percentage dry weight 92 16 Forage samples c o l l e c t e d i n the spring, 1966, i n regions 1, 2 and 3 at the Toby Creek l i c k , showing the sodium and potassium concentrations as a percentage dry weight 93 17 Chemical analyses of 3 l i c k s , comparing high and low l i c k i n g s i t e s 2.02 18 Some serum sodium values and the packed-cell volume obtained for the mountain goat 108 19 Selenium values for plant samples and earth l i c k samples from the P u r c e l l mountains, on a dry matter basis X LIST OF FIGURES To follow page 1, Outline map of B r i t i s h Columbia showing the location of the study area, i n the East Kootenay region 5 2, The i n i t i a t i o n and period of use at three low e l e v a t i o n a l l i c k s during the spring and summer of 1966 2M-3, The i n i t i a t i o n and period of use at three high e l e v a t i o n a l l i c k s during the spring and summer of 1966 2M-*f. The cumulative t o t a l s for 10 day periods, at the Toby Creek l i c k , showing d i f f e r e n t i a l use i n time 37 5, The increase i n female use of l i c k s during June, after the kidding period 38 6. The i n i t i a l date kids were observed at the Toby Creek l i c k and the increase i n kids at the l i c k during June 38 7» D i f f e r e n t i a l use i n time at the Dutch Creek l i c k using combined data from the summers of 1965 and 1966 39 8. D i f f e r e n t i a l use i n time at the Stoddart Creek l i c k during the summer of 1966 39 9. D i f f e r e n t i a l use i n time at the Lazy Lake l i c k during the summer of 1966 39 10. The average t o t a l group size ( a l l sex and age categories) per month, using observations from a l l l i c k s kh 11. The average group size for males (using a l l age categories) per month, using observations from a l l l i c k s h$ 12. The average group size for females (using a l l age categories) per month, using ob-servations from a l l l i c k s 1+6 To follow page 13. The sequence of molt as i t i s affected by-physiological processes and abrasion by vegetation while the animals are using the l i c k 5^ lk. The r e l a t i o n between environmental factors and animal numbers i n regions 2 and 3 during early spring of 1966 70 15o The r e l a t i o n between environmental factors and animal numbers i n regions 2 and 3 during June of 1966 70 16. The r e l a t i o n between environmental factors and animal numbers i n regions 2 and 3 during July of 1966 70 17. The r e l a t i o n between environmental factors and animal numbers i n regions 2 and 3 at the Dutch Creek l i c k during July of 1965 70 18. A comparison of forage samples c o l l e c t e d i n the spring showing sodium concentration , plotted against e l e v a t i o n a l region 9*+ 19. A comparison of forage samples c o l l e c t e d i n September showing sodium concentration plotted against e l e v a t i o n a l region 9^ 20. A comparison of forage samples c o l l e c t e d i n the spring showing ash as a percentage dry weight plotted against e l e v a t i o n a l region 9*+ 21. A comparison of forage samples c o l l e c t e d i n September showing ash as a percentage dry weight plotted against e l e v a t i o n a l region 9*+ 22. A comparison of spring and f a l l forage samples from the Toby Creek l i c k using sodium concentration as a percentage dry weight 95 23. A comparison of spring and f a l l forage samples from region 1 using sodium con-centration as a percentage dry weight 95 2h, A comparison of spring and f a l l forage samples from the Toby Creek l i c k using 95 x i i To follow page sodium concentration as a percentage dry-weight 95 25. A comparison of spring and f a l l forage samples from region 1 using sodium con-centration as a percentage dry weight 95 26, A high l i c k i n g s i t e i n the roots of a Douglas f i r tree at the Lazy Lake l i c k . A mouse trap marker i s shown i n the fore-ground 97 27. A high l i c k i n g s i t e i n a clay hank at the Mary Creek l i c k 97 x i i l ACKNOWLEDGEMENTS The f i e l d work of the project was financed by a National Research Council research grant, held by Dr. Ian McTaggart Cowan. The Divisions of Plant Science and S o i l Science i n the Faculty of Agriculture provided laboratory f a c i l i t i e s and able technicians to aid with a n a l y t i c a l work. The Department of Zoology provided o f f i c e space. Serum samples were analyzed by the laboratory technician of the Kimberley and D i s t r i c t H ospital. I am e s p e c i a l l y g r a t e f u l to Dr. Ian McTaggart Cowan, Dean of Graduate Studies, who suggested and directed the study. Dr. V.C. Brink, Chairman, D i v i s i o n of Plant Science, and Dr. H.D. Fisher, Dr. J.M. Taylor, Dr. H. Nordan of the Department of Zoology, provided valuable assistance and advice throughout the study. To my brothers, Don and Keith Hebert, who provided excellent f i e l d assistance and the rest of my family who gave time and support to the study, I am deeply indebted„ To t h e s e people, and the many others who contributed personal and t e c h n i c a l assistance, I would l i k e to extend my thanks„ INTRODUCTION 1 The Rocky Mountain goat (Qreamnos americanus B l a i n v i l l e ) ) occupies a r e l a t i v e l y wide l a t i t u d i n a l range i n western North America (M+ degrees to 61 degrees north l a t i t u d e ) . Within t h i s area i t s e c o l o g i c a l tolerance i s probably wider than any other large mammalo Goats are commonly found down to the edge of s a l t water on the coast of B r i t i s h Columbia where annual p r e c i p i t a t i o n exceeds 200 inches and on the a r i d slopes of Idaho where p r e c i p i t a t i o n may not reach 10 inches,, It i s a creature of steep t e r r a i n , where c l i f f s are a general physical feature and where vegetation i s sparse. In general, goats do not leave the steep mountain slopes where they can f i n d easy protection i n c l i f f s . A notable exception occurs during the summer i n the Rocky Mountains and Puree 11 Mountains when Qreamnos may leave the mountain slopes and t r a v e l through dense vegetation to reach exposed areas of mineral-rich earth l o c a l l y known as " l i c k s " . The l i c k s are seldom close to secure escape t e r r a i n or to good feeding grounds. It i s apparent, therefore, that i n seeking the l i c k s the animals are expressing a compulsion so great 1. S c i e n t i f i c names and a u t h o r i t i e s of mammals based on Cowan and Guiget (1965)« 2 that i t overrides the usually strong a f f i n i t y for alpine-type range. The compulsion draws the animals from available range and brings them by narrow, w e l l defined t r a i l s to areas more accessible to predators. This poses the idea that such behavior must r e f l e c t a need, the s a t i s f a c t i o n of which i s so important to the species that the possible a d d i t i o n a l r i s k s are tolerable at s the species levelo In t h i s region, almost a l l species of wild ungulates make use of mineralized earth l i c k s or mineral springs but the distance t r a v e l l e d and the degrees of departure from normal habitat and habits are greater i n the goat than i n any other species. The widespread occurrence of th i s habit among wild ungulates i n the d r i e r parts of western North America gives r i s e to a number of questions that require answers i f we are to understand the role of the l i c k s i n the ecology of species using them. What i s the nature of the demand f e l t by the animals 5 what mineral elements are they seeking from th i s s o i l and what part do they play i n the physiology of the animal; why i s i t necessary for them to seek these supplementary sources; to what extent does the presence of these l i c k s govern the h a b i t a b i l i t y of adjacent feeding grounds; how frequently and at what seasons must the animals have access to the l i c k s | how far w i l l they come to reach them; what 3 a d d i t i o n a l mortality from predation r e s u l t s from the habit of using l i c k s ? There have been attempts to answer some of these questions for several of the lic k - u s i n g species. In general, studies have concentrated upon attempts to determine a sought-after mineral or minerals. Thus Cowan and Brink (19^9) examined samples taken from 11 l i c k s i n the Rocky Mountain National Parks of Canada i n the hope that chemical analysis would reveal the needed element. Other workers, notably Honess and Frost (19^2), Stockstad (1953) and Williams (1962) have used a similar approach. A second method of attacking the question has been to es t a b l i s h mineral " c a f e t e r i a s " wherein a wide var i e t y of elements i n d i f f e r e n t formulations have been exposed to open s e l e c t i o n by the wild species (Staith, 195^; Stockstad, 1953 and B i s s e l , 1953). The present study was undertaken to gain a d d i t i o n a l information on the use of l i c k s by Qreamnos. I wished to document some of the factors a f f e c t i n g the patterns and nature of use by the d i f f e r e n t age and sex categories, the frequency of use by the same i n d i v i d u a l , l i c k structure and how i t affects patterns of use and the distance t r a v e l l e d to reach the l i c k s . An attempt was to be made also, to gain further insight into the nature of the demand that the l i c k s were s a t i s f y i n g and to determine whether there was a detectable mineral deficiency i n the animals associated with d e f i c i e n c i e s i n the normal food plants 0 Also to be explored were the a d d i t i o n a l r i s k s en-countered when the animals l e f t their alpine ranges and their impact on the animals using the l i c k 0 5 DESCRIPTION OF THE AREA The study area i s centered i n the Rocky Mountain Trench between Cranbrook and Radium Junction (H-9°30s to 50° north latitude) as shown i n Figure 1 0 The Trench i s a l o n g i t u d i n a l , glaciated depression extending from Montana to the Yukon. The Rocky Mountains r i s e abruptly from the v a l l e y f l o o r on the east and the Puree11 Mountains on the west begin as rounded and wooded f o o t h i l l s , which give way to rugged mountains (Holland, 196*+) „ The climate i s such that there.is higher p r e c i p i -t a t i o n i n winter than i n summer and a high proportion &f snowfall. It i s semi-arid with a range of annual p r e c i p i t a t i o n of about lk to 17 inches 0 The heaviest f a l l s of r a i n and snow are on the western slopes (Kelley and Holland, 1961). The area i s characterized by four major vegetation zoness the Ponderosa pine-bunchgrass zone with i t s l i g h t and dark brown s o i l s occurs at an elevation of approximately 2500 feet; the Interior Douglas f i r zone has the orthic brown s o i l s at an elevation of 2000 to k$00 feet; the Engelmann spruce-subalpine f i r zone has o r s t e i n podzol s o i l s at an elevation of 3500 to 5500 f e e t | the alpine zone with i t s t h i n humus s o i l s occurs around 7000 feet (Krajina, 1965).. The aspect of the slope w i l l a f f e c t the e l e v a t i o n a l l i m i t s of each zone, to some degree,. FIGURE 1. Outline Map of British ColuMbia showing the location off the study area,in the East Kootenay region. 6 During the early spring, Oreatnnos leaves the winter ranges and passes through the alpine and spruce-fir zones to reach l i c k s i n the Douglas f i r zone. The physiography of the Douglas F i r zone i s such that the animals u t i l i z e available c l i f f s for food, escape t e r r a i n and as a base from which they t r a v e l to the l i c k s . In essence, each l i c k i s an accumulation s i t e located on a bench or l e v e l l i n g off of a slope,, Runoff, depending on the grade of the slope above the l i c k , c a r r i e s minerals from higher vegetation zones to the l i c k region where they are deposited. The l i c k s i n the study area w i l l d i f f e r i n degree of accumulation of minerals since the Stoddart Creek, Mary Creek, E l k Creek, and Lazy Lake l i c k s are situated i n the Rocky Mountains while the Toby Creek, Dutch Creek, and Findlay Creek l i c k s l i e i n the Puree11 Mountains and these two ranges d i f f e r i n topography and r a i n f a l l . Although a l l l i c k s are i n the Interior Douglas f i r zone e l e v a t i o n a l differences among l i c k s i t e s cause some l i c k s to-border the spruce~fir zone and others the bunchgrass zone« The Stoddart Creek l i c k at an elevation of 2900 feet i s on a south facing slope which borders the open park-land of Douglas fir-bluebunch wheatgrass. The stretches of grassland on dr i e r slopes contain bluebunch wheatgrass 7 (Agropyron spicatum). Idaho fescue (Festuca idahoensis)„ and June grass (Koelerla c r i s t a t a ) and serve as sheep (Q-ffls canadensis) winter range since snowfall i s usually low. The Toby Creek l i c k on the north-facing bank of the creek i s at an elevation of 3 0 0 0 f e e t . It i s centered i n the Douglas f i r - p i n e grass subzone of the wooded f o o t h i l l s of the Puree l i s which may to some extent serve as moose (Alces alces) winter range. The Dutch Creek and Findlay Creek l i c k s border the spruce-fir zone at an elevation of about ^000 f e e t . They are both on west facing slopes i n an area of high snowfall. The E l k Creek and Mary Creek l i c k s i n the Rockies, border the spruce-fir zone at an elevation around k$00 f e e t . Both are found on southwest facing slopes where elk (Cervus  canadensis) migrations are common. The Lazy Lake l i c k i s situated on a southwest facing slope at an elevation of hooo f e e t . It borders the Ponderosa pine-bunchgrass zone since the Rockies r i s e abruptly i n t h i s region and elevation becomes less important i n separating zones. The area i s extremely dry with a l i g h t snowfall i n winter. The behaviour of the mountain goat during the spring and summer leads i t to occupy three e l e v a t i o n a l regions for varying lengths of time. As the winter snow recedes goats move into the alpine and subalpine zones which serve as summer 8 range for approximately six months. Animals leave t h i s range for short periods to t r a v e l to natural earth l i c k s to acquire mineralso While i n the v i c i n i t y of the l i c k they occupy a region which i s e s s e n t i a l l y a modified form of the i r summer range. It offers escape t e r r a i n i n the form of c l i f f s , food, cover and bed s i t e s . In general, t h i s region i s about 1500 to 2000 feet higher i n elevation than the respective l i c k , while the summer range occurs between 6000 and 8000 f e e t . These three regions may be designated 1, 2, and 3 and occur to a greater or lesser degree at each l i c k studied. For any of the l i c k s studied, region 1 i s the summer range the l i c k serves; region 2 i s the temporary safety area close to the l i c k used while the animal i s preparing to v i s i t the l i c k proper, or between several v i s i t s to the l i c k In quick succession; region 3 i s the l i c k i t s e l f . 9 METHODS AND MATERIALS The methods used during t h i s study consisted of f i e l d methods to determine patterns of use and laboratory methods to determine mineral content and to detect mineral d e f i c i e n c i e s of the vegetation, animals and l i c k s o i l s . Ob-servations i n the f i e l d were designed to show, at the l i c k s , frequency of use by the same i n d i v i d u a l , d i f f e r e n t i a l use i n time by the sexes, movement between designated areas and the r e l a t i o n of the factors a f f e c t i n g these patterns. The lab-oratory methods compared mineral content of forage species during the spring, summer and f a l l seasons at d i f f e r e n t elevations and mineral content of the blood before and after l i c k use to see i f d e f i c i e n c i e s could be detected. Lick samples were analyzed to see i f the mineral content was s u f f i c i e n t to supplement d e f i c i e n c i e s i n the plants. T r a i n s The two main purposes of trapping were to obtain blood serum samples and to mark animals so that information could be obtained regarding patterns of movement. Two types of traps were used during the study. Permanent c o r r a l traps b u i l t on well defined t r a i l s had a gate on one or both ends and were operated by a trigger rope. They were approximately 8 feet high, 20 feet long and the width varied depending on 10 the t r a i l . Each was lined with chicken wire i n case poles came loose during any struggle by the animalo The trap at Dutch Creek had two gates while those at Toby Creek and Findlay Creek had only one 0 Two moveable box-type traps were obtained from the B„Co F i s h and.Wildlife Service for use during the summer,. They were approximately 5 feet long, h feet wide and h feet high. They had a gate at one end, were covered with ^ inch thick, 2 inch rope mesh and had a frame made of *jr inch black pipe. Marking Trapped animals were marked by painting their horns with coloured enamel from spray cans. A d i f f e r e n t color was used for each animal at any one pa r t i c u l a r l i c k . Also, a number approximately 1„5 to 2 feet high was painted on both sides of each animalo Animals were marked without trapping using two types of automatic marking devices„ The simpler device con-s i s t e d of a rat trap, an egg f i l l e d with h i s t o l o g i c a l dye and a trigger stringo The device was assembled on a s t r i p of plywood h inches wide and 6 feet long and suspended above a t r a i l entering the l i c k . Twenty of these devices were used during the summer. The eggs were f i l l e d with h i s t o l o g i c a l dye using a 250 m i l l i l i t r e veterinary syringe. A t r i p s t r i n g was made of 25 pound test nylon f i s h l i n e . A piece of wire 11 was attached to the snap wire of the trap, to cut the egg cleanly, allowing the dye to drop d i r e c t l y down onto the animal,, Dye colors were rotated i n each device during the summero The automatic spray device was adapted from Clover (1955)o It consisted of a pressure tank containing dye and 120 pounds a i r pressure, a nozzle to di r e c t the flow and a pressure treadle triggered by the animal to release the dye. The method used to prepare the dyes was taken from Hansen (1963), The dyes used were rhodamine B extra, malachite green c r y s t a l s and p i c r i c a c i d . These gave pink, green and yellow c o l o r s , respectively„ Green and pink solutions were mixed to give v i o l e t . The pink and green dyes were prepared by diss o l v i n g rhodamine B extra or malachite green, res-pectively, i n 1 pint of a 99 percent solution of isopropyl alcohol,, To thi s an equal amount of water was added. The p i c r i c acid was made into a saturated sol u t i o n with isopropyl alcohol and an equal amount of water added. Vegetation Before forage species were c o l l e c t e d to determine mineral content, rumen samples from 5 goats shot i n the study area were examined to get a r e l a t i v e idea of the food types. Samples were preserved i n 10 percent formalin. The material used for examination was strained, washed and food types i n 12 the form of grasses and sedges, forlDS, shrubs and conifers were separated. The displacement technique, employing a graduated cylinder f i l l e d with water was used to determine the volume of each 0 From t h i s , each food type was calculated as a percentage of the t o t a l . Animals feeding i n the v i c i n i t y of the l i c k s were observed to see which forage species were being u t i l i z e d . Forage species were c o l l e c t e d from May u n t i l September during the summer of 1966 with an attempt being made to c o l l e c t plants i n the same phenological growth stage. The actual c o l l e c t i n g consisted of taking 10 to 20 annual-growth stems (Including leaves) from as many i n d i v i d u a l plants as possible, for each species. Plants c o l l e c t e d i n late May (spring c o l l e c t i o n ) were taken from the v i c i n i t y of the l i c k . In early June plants were c o l l e c t e d from region 2. In late June and early July plants were c o l l e c t e d from region 1, Since receding snow cover l i m i t s spring growth, plants at higher elevations began to grow and mature l a t e r . Therefore, plants c o l l e c t e d i n region 1 should have been at approximately the same growth stage as plants c o l l e c t e d at the l i c k . During late August and early September the same species were again c o l l e c t e d at the same e l e v a t i o n a l s i t e s . Plants at a l l s i t e s were completely mature with seed^heads 13 and berries abundant. Although plants c o l l e c t e d at the l i c k had been i n the maturation stage longer than plants c o l l e c t e d i n region 1 t h i s factor was not expected to a l t e r the r e s u l t s , as p r e c i p i t a t i o n and consequent leaching had been n e g l i g i b l e c New-growth stems-of the browse species involved were removed at the nodes and grass was clipped just above the base (Dietz et. a l . 1962)„ Each species was placed i n a paper sack marked with the date, species, l i c k and e l e v a t i o n a l s i t e . A l l samples were a i r dried during the summer0 Larger and coarser samples were ground i n a hammer m i l l ; smaller samples were ground i n a Wiley m i l l . The ground material was mixed and. placed i n glass bottles u n t i l needed for subsequent chemical analysis. As far as possible a l l plants were c o l l e c t e d at Toby Creeko In some cases, plants from regions 1, 2 or 3 at other l i c k s were used when the par t i c u l a r species could not be obtained at Toby Creeko Since l i c k s are accumulation s i t e s , It was supposed that.plants c o l l e c t e d at the same el e v a t i o n a l regions__at_Qthe.^>licksywould.,he under approximately the same co.nditlons.vof leaching* landimovement of minerals down the slope 0 . .Analysis for sodium and potassium i n the plant was done as described by Chapman and Pratt ( 1 9 6 l ) a Approximately 3 grams of each sample were placed In a crucible and- dry ashed at 600 degrees fahrenheit 0 The ashed material was Ih dissolved i n 20 percent HCl and di l u t e d to 100 m i l l i l i t r e s with d i s t i l l e d water. The contents were placed i n flasks and stoppered u n t i l needed. Standard curves for sodium and potassium were plotted £or the Perkin-Elmer flame photometer and the unknown solutions run.accordingly. Concentrations as percent dry matter i n the plant were calculated from the curves and are shown i n Tables 15 and 16. Lick S o i l s Lick samples were c o l l e c t e d from the Dutch Creek, Toby Creek, Mary Creek and Lazy Lake l i c k s so that two types of samples were taken from each l i c k . Observations showed that l i c k s are composed of h i g h - l i c k i n g s i t e s and low-licking s i t e s which may or may not be adjacent to one another. Surface material was c o l l e c t e d from each s i t e so that i t represented what the animals were actually l i c k i n g or refusing. The material was put through a screen, a i r dried and stored i n glass j a r s . S o i l samples were analyzed using the method adopted by Stockstad (1953)» described below. Chemical analysis was ca r r i e d out on 11 samples from 3 l i c k s 0 A l l s o i l samples were treated with ammonium acetate at two pH l e v e l s 0 The amount of each element available to the animals using the l i c k s was approximated by using buffered extracting solutions with pH values similar to those found i n a ruminating animal's 0 15 abomasum and intestine„ A series of pH determinations on the digestive tracts of 30 elk and deer indicated that pH values of *+.00 and 7*00 should be used. Phosphorus con-centrations were determined using the Bray method for extraction purposes. The actual determinations, for a l l elements, were made by following the procedures outlined by the Association of O f f i c i a l A g r i c u l t u r a l Chemists (I960), Blood Blood samples were taken i n the f i e l d using vacu-tainer tubes and needles 0 The vacutainer tubes drew approx-imately 10 m i l l i l i t r e s and contained no anticoagulant. The needles were 1,5 inches i n length. A l l samples were taken from the jugular vein. C a p i l l a r y tubes containing heparin as the anticoagulant were used to obtain hematocrit values. Test tubes containing heparin were substituted i n June for the c a p i l l a r y tubes due to loss and breakage„ Blood samples kept c o o l i n an insulated cooler were transferred to the cooler i n the Kimberley Ho s p i t a l within two hours after the sample was taken. The serum was separated on a centrifuge and analyzed for sodium using a flame photo-meter with an error of plus or minus two percent. The hep-ari n i z e d samples were centrifuged and the packed-cell volume determined as a percentage, from a known volume of blood. A l l analyses were done by the laboratory technician i n the Kimberley and D i s t r i c t H o s p i t a l . 16 RESULTS L ICK STRUCTURE The d r y e a r t h l i c k s a r e f o r m e d n a t u r a l l y and t h e g r e a t v a r i a t i o n i n s i z e and s t r u c t u r e among them c a u s e s d i f f -e r e n c e s i n p a t t e r n s o f u s e . E a c h l i c k i s l o c a t e d In an a c c u m u l a t i o n s i t e and has a t r a i l s y s t e m b e t w e e n r e g i o n s 1, 2 and 3 t h a t i s s i m i l a r i n b a s i c d e s i g n . The l i c k s d i f f e r i n s i z e , s t r u c t u r e and c o m -p l e x i t y . W i t h i n t h e l i c k , t h e a c t u a l l i c k i n g s i t e may be a s m a l l m o i s t p o c k e t o f c l a y , a c a v i t y b e n e a t h a D o u g l a s f i r t r e e o r a c o m b i n a t i o n o f t h e t w o D T r a i l s y s t e m s The s t r u c t u r e o f t h e t r a i l s y s t e m i s s u c h t h a t e a c h l i c k I s c o n n e c t e d by a w e l l d e f i n e d t r a i l t o r e g i o n 2 and r e g i o n 2 has a n e t w o r k o f t r a i l s b r a n c h i n g f r o m one m a i n t r a i l w h i c h l e a d s i n t o a l p i n e summer r a n g e . E a c h l i c k i s b o u n d e d on 3 s i d e s by w e l l d e f i n e d t r a i l s s o t h a t g o a t s c a n e n t e r o r l e a v e t h e l i c k o n two s i d e s and j o i n w i t h t h e t r a i l f r o m r e g i o n 2 on t h e t h i r d s i d e c The f o u r t h s i d e u s u a l l y ha s a r i v e r o r c r e e k f o r m i n g I t s b o u n d a r y . The t r a i l b e t w e e n r e g i o n 2 and t h e r e s p e c t i v e l i c k p a s s e s t h r o u g h d e n s e t i m b e r , d o e s n o t b r a n c h o r s p l i t and i s a b o u t | t o 1 m i l e l o n g . T h e t r a i l f r o m r e g i o n 1 t o r e g i o n 2 p a s s e s t h r o u g h t h e D o u g l a s 17 f i r zone, the spruce-fir zone and into the alpine zone. It may be 5 to 15 miles long and follows ridges and skree slopes. A description of the use of t r a i l systems i s needed to show the s p e c i f i c i t y of movement from one region to another and the degree of selectiveness goats show between major and minor l i c k s * In the early spring, the males move from th e i r winter ranges to region 2 where they begin feeding on succulent forage. From region 2 they make frequent t r i p s to the l i c k and for about two weeks can be found i n the l i c k or region 2. They then move onto th e i r respective summer ranges 0 At no time during the spring or summer was shed hair or f e c a l material found anywhere but on the t r a i l system between the three regions. Goats did not appear to wander, as deer do, but used the t r a i l s to reduce t r a v e l l i n g time between regions. This made them less vulnerable to predators and they spent less time i n habitats low i n palatable forage. At the Toby Creek l i c k , on two occasions, goats were found wandering down a road which crosses the t r a i l between region 2 and the l i c k . It appeared as i f the goats had missed the t r a i l while crossing the road and, as they moved down the road, had ex-amined each opening Into the brush. As none of the openings were connected to the t r a i l they kept moving down the road. In both instances they moved back up the road and eventually found the t r a i l . In neither case did they move through the 18 timber to reach region 2, Kids, when separated from their mothers i n the v i c i n i t y of the l i c k always returned to the t r a i l leading to region 2 rather than move through the timber. More wandering occurred between region 2 and region 1 when the t r a i l was above timberline. This was attributed to the animals moving to i n d i v i d u a l summer ranges and to the increase i n palatable forage species. The movement of goats between regions i s very def-i n i t e and the t r a i l system i s used to increase e f f i c i e n c y . In much the same way, goats are very e f f i c i e n t and selective i n the l i c k s they use. At the Dutch Creek, Lazy Lake and Stoddart Creek l i c k s a major and.a minor l i c k were present. In a l l 3 instances, the minor l i c k was more accessible and closer to region 2. The goats would move past minor l i c k s and u t i l i z e a major l i c k . Use of the minor l i c k Increased at the Dutch Creek and Stoddart Creek l i c k s over the summer but thi s was probably caused by trapping operations at the major l i c k . The goats were very sel e c t i v e i n the l i c k they chose and also very selective In the actual l i c k i n g s i t e within the l i c k . At Lazy Lake, the t r a i l passed through a minor l i c k north of the major l i c k yet l i t t l e or no l i c k i n g occurred except i n the major l i c k . The c a v i t i e s i n the minor l i c k appeared older and contained more rock. The minor l i c k s were smaller than the major l i c k s i n a l l Instances, 19 Size and structure The l i c k s d i f f e r so greatly i n size and structure that a detailed description of each i s not possible. The Toby Creek l i c k i s about 2 miles long and 200 yards wide; the Dutch Creek l i c k i s about 100 yards by 200 yards; the Lazy Lake l i c k i s about 300 yards by *f00 yards and the Stoddart Creek l i c k i s about 100 feet by 200 feet. The Stoddart Creek and Dutch Creek l i c k s are open clay pits pro-viding no cover or escape t e r r a i n . They are very exposed and the hot summer sun of July and August keeps the clay hot and dry. The Lazy Lake l i c k has a l l l i c k i n g s i t e s among the roots of Douglas f i r trees. The Toby Creek l i c k has s t r i p s of timber, 3 small clay pits among the roots of Douglas f i r trees i n a timbered section, escape t e r r a i n i n the form of steep c l i f f s and exposed clay banks. The complexity of the l i c k i s determined by comparing the habitat features of region 1 to those found i n the l i c k . The Toby Creek l i c k i s the most complex since i t contains almost a l l the habitat features found i n region 1. The' Lazy Lake l i c k contains timber, some palatable forage and some escape t e r r a i n . The Stoddart Creek and Dutch Creek l i c k s are the least complex, being only ex-posed pits having no other habitat features. The Dutch Creek l i c k contained some palatable forage around i t s edge. PATTERNS OF MOVEMENT AND LICK USE 20 Introduction The actual s a l t requirements of big game have never been extensively investigated and no knowledge of a n u t r i t i o n a l necessity i s av a i l a b l e . Inasmuch as i t has been conclusively demonstrated that both sodium and chlorine are e s s e n t i a l for most animal l i f e , a phy s i o l o g i c a l necessity existing i n wild ruminants would be no exception. Since sodium i s not con-sidered necessary for normal growth of most plants, i t i s highly debatable whether a c e r t a i n range can f u l l y s a t i s f y the ph y s i o l o g i c a l requirements of sodium i n the animal. The chemical nature of the range would to a great degree determine the n u t r i t i o n a l requirement of s a l t i n ungulates. The importance of the patterns of movement exhibited by the mountain goat i s seen when we r e a l i z e that they place the animal on ranges varying i n chemical composition. They also serve to determine the degree of contact with other species, parasites and predators. It i s necessary that these patterns be described and understood i n order to assess the vegetative conditions an animal encounters prior to entering the l i c k . In order to show that the diet of the goat can not supply s u f f i c i e n t sodium, the feeding patterns i n r e l a t i o n to the mineral content of the forage must be correlated. It has been suggested that ungulates use l i c k s i n 21 an attempt to supplement a di e t low i n mineral content 0 This leads to the conclusion that there are many minerals which could be d e f i c i e n t i n the vegetation and which could attr a c t the animals to natural l i c k s e Evidence i s being accumulated to show that spring forage conditions may play an important role i n i n i t i a t i n g l i c k use 0 Dalke et, a l , (1965) found that a conspicuous s a l t hunger i n elk developed 2=3 weeks after early use of succulent forage. It would seem that a winter diet low i n sodium, would, upon eating spring forage high i n water con-tent with l i t t l e or no sodium, Induce a physiological state i n the animal whereby sodium was d e f i c i e n t . It has been found by Marriott (1950) that "pure s a l t depletion" occurs i n patients who become d e f i c i e n t i n sodium and chlorine while being l i b e r a l l y supplied with water, A patient on a s a l t d e f i c i e n t diet lost 5»96 grams of sodium i n the f i r s t 10 days but after that the kidney, i n i t s regulatory function, greatly reduced the sodium output. A l l studies with mineral cafeterias have, .shown..conclusively that sodium was the im-portant e_lament. sought after under the prevailing conditions (Stockstad, 19535 B i s s e l , ,1953 and Smith, 195^)° The present study w i l l attempt to relate patterns of movement of the goat to the sodium concentration of the forage, the l i c k s o i l s and the blood serum of the animals. 22 Seasonal patterns In the Rocky Mountain Trench of B r i t i s h Columbia natural mineral l i c k s are abundant and widely d i s t r i b u t e d . They appear a t t r a c t i v e to large populations of wild ungulates, although l i t t l e i s known about the patterns of use by i n -d i v i d u a l species. Licks situated at the lower edge of the Douglas f i r zone are. adjacent to winter feeding grounds of bighorn (Ovis canadensis)« moose (Alces a l c e s ) , elk (Cervus  canadensis) and mule deer (Odocoileus hemionus). Licks at higher elevations l i e on the migration routes of these species and are u t i l i z e d during the spring migration. In general, mountain goats winter at high elevations on exposed wind-swept slopes or adjacent timber and c l i f f s , while the above mentioned species winter i n the bunchgrass zone or adjacent timber. This trend reverses somewhat i n the summer; Individual goats t r a v e l to natural l i c k s at low elevations while other species move to alpine regions. The s h i f t occurs i n May and June as the snow pack recedes. The big game species mentioned, with the exception of the goat, move into the v i c i n i t y of the l i c k s i n late f a l l (November) and remain there u n t i l late spring (June), having access to the l i c k s for approximately 8 months. The goat uses l i c k s from early spring ( A p r i l ) to early f a l l (September). Dalke ejj. ajlo (1965) states that use of s a l t l i c k s by elk was n i l during February and March, and there was 23 no evidence that elk pawed through the snow to gain access to the s a l t . Deer, sheep and caribou have been observed l i c k i n g s a l t i n the winter, on roadsides, where man has spread i t to improve road conditions 0 There i s no evidence to suggest that goats use natural or a r t i f i c i a l s a l t l i c k s at any time during the winter months„ The use of l i c k s In t h i s area, by ungulates, pro-duces a concentration of big game species i n the v i c i n i t y of the l i c k during the l a t t e r part of A p r i l , a l l of May and i n i t i a l part of June, Mineral springs are rendered unavailable In winter by freezing but clay from earth l i c k s i s probably available since material i s chewedo This leads to the con-clus i o n that seasonal concentrations of ungulates are based upon factors other than those of a v a i l a b i l i t y of l i c k s o i l s . A r t i f i c i a l s a l t l i c k s were used by elk on their migration only as the upper edge of a c t i v i t y reached them. No pawing down i n the snow to reach s a l t l i c k s ahead of time was noted, while elk were seen to dig as deep as h-8 inches i n snow for c e r t a i n desirable food plants„ The apparent centre of elk a c t i v i t y was well below the l i c k s for some time after they had become snow free (Beeman, 1957)o The i n i t i a t i o n of l i c k use by mountain goats would seem to be an extremely variable factor„ It appears to be affected by elevation and receding snowpack, spring vegetation 2h and weather. The f i r s t day of use i s recorded for each l i c k i n Figures 2 and 3« Figures are for the summer of 1966 since work was not started u n t i l June i n 1965. I n i t i a l use was mainly by males according to observations made i n early spring. The numbers were very low during early May but gradually In-creased u n t i l a peak was reached i n June. The majority of goats were observed feeding rather than l i c k i n g upon descent to low elevations. This suggests that i n i t i a t i o n of l i c k use more accurately describes movement into the v i c i n i t y of the l i c k . Termination of use at each l i c k varies i n r e l a t i o n to the t o t a l period of use from the i n i t i a t i o n of l i c k use. High e l e v a t i o n a l l i c k s such as Dutch Creek and E l k Creek had goats using them on August 15, 16 and 23| with probable use to the end of August or early September. Records of goats shot In the Dutch Creek l i c k i n September of previous years attest to t h i s statement. Goats using low e l e v a t i o n a l l i c k s such as Toby Creek and Stoddart Creek terminate l i c k use around the end of July. Only one female and kid were seen i n the Toby Creek l i c k at t h i s time and only one lone adult at the Stoddart Creek l i c k . After t h i s , no tracks or sign of animals using the t r a i l could be found. In general, low e l e v a t i o n a l l i c k s are used early In the spring with termination of use occurring i n late summer. STODDART CREEK TOBY CREEK LAZY LAKE 30 123456789 10 April May T I M E 20 21 22 23 24 25 26 27 28 29 30 31 12 3 4 5 6 7 July August F I G U R E 2. The initiation and period of use at three low elevational licks during the spring and summer of 1966 MARY CREEK FINDLAY CREEK DUTCH CREEK 10 12141618 20 22 24 26 28 30 1 3 5 7 9 May June 1517192123 25 2729 31 24 6 8 August September TIME FIGURE 3. The initiation and period of use at three high elevational licks during the spring and summer of 1966 25 High e l e v a t i o n a l l i c k s are not used u n t i l late spring but use continues l a t e r into the f a l l . The t o t a l period of use appears to be similar for these two groupings of l i c k s . E l e v a t i o n appears to be an important factor reg-ulating i n i t i a t i o n of use and t o t a l period of use„ In general, the l i c k s at low elevations, used during A p r i l and May have a reduced snow cover and the surrounding area supports early spring growth. The Lazy Lake l i c k probably receives the least snow and has the e a r l i e s t growth of vegetation, since i t i s on an exposed southwest facing slope bordering the bunchgrass zone. The snow recedes by late A p r i l , exposing the whole face, and spring growth begins i n May, Goats observed at t h i s time are at elevations approaching that of the l i c k or lower. In most cases they were seen feeding on a low c l i f f (at 3000 feet) which r i s e s abruptly from the v a l l e y f l o o r . At Toby Creek and Stoddart Creek, i n i t i a l use was somewhat l a t e r . Region 2 at both l i c k s i s a north facing slope which retains Its winter snow pack. Males, f i r s t to a r r i v e , fed on the lower edge of the c l i f f s , after spending much time i n small openings i n the timber below the c l i f f s . The high e l e v a t i o n a l l i c k s such as Dutch Creek and Mary Creek r e t a i n their winter snow longer and are more susceptible to spring snow storms since region 2 occurs 26 between 5000 and 6000 f e e t . In May, at the Mary Creek l i c k , region 2 was covered frequently by spring snow storms. Two sets of c l i f f s are spaced one above the other, above the l i c k , one about 1000 feet higher than the other. The top set con-tained winter snow into late May while the lower set was often covered with snow from spring storms. At the Toby Creek and Lazy Lake l i c k s a d e f i n i t e pattern of feeding existed from i n i t i a l use i n the spring to termination i n the f a l l . There was a continual increase In the elevation of feeding s i t e s over the summer. In early spring, the goats at Lazy Lake fed at the elevation of the l i c k or lower, and the goats at Toby Creek fed on the lowest c l i f f s . By the end of May or early part of June the animals at Lazy Lake were ranging i n the region of the pinnacle and c l i f f s around i t (approximately 1500 to 2000 feet higher than the l i c k ) . At Toby Creek the animals were using the lower c l i f f s on the west side of region 2, By the end of June or early part of July when the females began moving i n , feeding occurred, on, the upper portion of the c l i f f s . By the end of July mo^sj^anima-ls^e^ on .the higher c l i f f s on the east side of region 2. Very, few .animals? wer.e. seen on the lower c l i f f s and those that were seen were usually just.passing through. At Lazy Lake, at th i s time, the same trends-occurred with fewer animals seen around the pinnacle and more moving 2? i n from region 1 d a i l y 0 Succulent spring vegetation as a determinant of seasonal patterns It has been shown that goats use natural l i c k s only i n the summer, that feeding on succulent vegetation usually precedes l i c k use and that goats feed at progressively higher elevations during the summer and are probably feeding on plants i n an early stage of growth for most of the summer0 according to Table 3> at least 87 percent of the goats ob-served at the Lazy Lake and Dutch Creek l i c k s were not i n the l i c k during May0 In A p r i l ? a l l goats were observed feeding and none l i c k i n g , In June, 72 percent of the observations showed goats l i c k i n g and 28 percent of the observations showed them feeding 0 Recent studies have shown t h a t u t i l i z a t i o n of succulent forage In the spring may contribute to i n i t i a t i o n of l i c k use 0 Dalke et„ ale, ( I 9 6 5 ) found that prior to the middle of May, the appetite of elk for new vegetation was greater than the desire for salt and that s a l t grounds within th© succulent herbaceous areas were used considerably more than the l i c k s i n shrubby areas. He also found that the ex-tent to which each of the natural l i c k s was u t i l i z e d by elk during any one period of the spring and summer depended c h i e f l y upon the elevation of the l i c k and i t s location with 28 respect to the various elk populations and the more preferred secluded areas. Williams (I962) found that the advent of green vegetation at the lower elevations attracted and held the animals i n the lower areas during a period from about mid A p r i l to the f i r s t of May. The elk migration or upward move-ment, once commenced, proceeded concurrently with the progress of forb, sedge and grass development and i t was during the gradual upward movement that concentration on s a l t grounds f i r s t became prominent. This movement of elk due to spring growth i s similar to that found i n goats at both the Toby Creek and Lazy Lake l i c k s . The fact that use of succulent vegetation precedes l i c k use may indicate a factor determining both the seasonal pattern of movement and the i n i t i a t i o n of l i c k use. It would suggest that the kidney can compensate for a winter diet low i n sodium but cannot maintain homeostasis i n the spring when ungulates increase their water intake on a low sodium d i e t . According to Kerr (1965) the downward movement of goats i n spring i s probably a response to new vegetative growth which begins to appear on the lower mountain slopes at that time. There appeared to be a corre-l a t i o n between the elevation at which animals were found and the new vegetative growth. As described, big game species winter at low e l e -vations i n th i s area and so take advantage of succulent spring 29 growth. They have a short distance to t r a v e l to the l i c k while the goat must t r a v e l long distances to c a p i t a l i z e on succulent vegetation and l i c k s . K l e i n (1953) suggests that the lack of available forage and shelter i n the high summer ranges during winter months causes f a l l movements from the alpine summer ranges to the subalpine f i r f o r e s t s . The spring d i s p e r s a l from the winter to summer range i s governed by the rate of upward retreat of the snowline. Wintering bands of goats move to the upper fringe of timberline with the f i r s t warm days of spring and remain Ih t h i s area u n t i l the snow cover has l e f t the higher treeless slopes 0 It appears then, that the pattern of feeding i n the spring and seasonal use of l i c k s by big game could be de-termined by the water and mineral content of spring vegetation. Seasonal patterns of movement invoke a d d i t i o n a l r i s k s Since the density of big game species increases i n the v i c i n i t y of the l i c k s i n early spring, i t was thought that the presence of other ungulate species might a f f e c t the seasonal use of l i c k s by goats. In t h i s sense, joint use among ungulate species was examined. Summer use of l i c k s also affects the goat adversely by bringing i t into contact with large populations of the paralysis t i c k (Dermdcentor andersoni). At t h i s time, goats encounter a greater abundance of predators and hunting pressure i n the f a l l . This suggests that the 30 benefits derived from l i c k use outweigh the sum of the negative factors and leads to the perpetuation of the species. The ef f e c t of each negative factor does not af f e c t each sex to the same extent and w i l l be explained from a viewpoint of d i f f e r e n t i a l use i n time. I n t e r s p e c i f i c joint use controlled by seasonal patterns V i r t u a l l y no i n t e r s p e c i f i c competition for l i c k use was observed at the various l i c k s . On some occasions other species besides goat were seen i n the l i c k . Almost no other tracks, shed hair or f e c a l material was found i n the l i c k . Moose and whitetailed deer had home ranges i n the v i c i n i t y of the Toby Creek l i c k but neither species was observed i n the l i c k . In early June, both elk and goats were observed at the Mary Creek l i c k . At thi s time elk were migrating from their winter to summer ranges and many paused at this high e l e v a t i o n a l l i c k which i s at the lowest point of elk summer range. Intensive l i c k use by elk occurred i n the early morning, late evening and at night. Goats used the l i c k mainly i n the afternoon. L i t t l e use was observed i n the morning or evening. S p e c i f i c l i c k s i t e s were found i n the bank at two d i f f e r e n t heights; those made by elk , being two to three feet higher than those made by the goat. Most elk were observed l i c k i n g on the east side of the l i c k and most 31 goats on the west side. E lk use was most intensive during the l a s t week of May and early part of June when goat use was just beginning and least intensive. It appears that joint use by two ungulate species existed at thi s l i c k for a short period but i n t e r s p e c i f i c competition was ne g l i g i b l e due to behavioural differences. Since the elk migration i s timed to follow the growth of spring forage, i t would appear that this i s a major factor bringing both species to the l i c k at approx-imately the same time. Goats, sheep and mule deer frequent the Lazy Lake l i c k during May and June. Both sheep and deer were observed using the l i c k and droppings and shed hair attested to their presence. Goats were never observed i n the l i c k at the same time as sheep or deer. The majority of goats were observed feeding on a low c l i f f north of the l i c k i n A p r i l and May, and i n June most sheep and deer migrated to summer ranges. Although joint use existed i t does not appear that competition reduced available l i c k use to goats. No evidence of competition or joint use was observed at the Dutch Creek l i c k . Mule deer, elk and moose were seen feeding i n the v i c i n i t y of the l i c k , but no tracks, shed hair or f e c a l material from these species were found i n the l i c k . At Stoddart Creek the l i c k i s about one mile from 32 sheep winter range. Local counts show that about 100 sheep are present over the winter. At no time were sheep seen i n the l i c k ; sheep tracks, f e c a l material or shed hair were not found i n the v i c i n i t y of the l i c k s . Sheep migrate to summer ranges i n Kootenay National Park where l i c k s are abundant and t h i s may be the reason joint use did not occur at the Stoddart Creek l i c k . Also, this l i c k i s i n the Douglas f i r zone which provides l i t t l e available forage. Goats u t i l i z e forage on nearby c l i f f s but sheep do not appear to t r a v e l through .the timber between the bunchgrass zone and the l i c k . It does not appear that competition i s important i n l i m i t i n g l i c k use by goats but joint use by other species Is common. Behavioural t r a i t s common to the species keep each from i n f r i n g i n g on the other's a c t i v i t i e s . Joint use occurred at high and low el e v a t i o n a l l i c k s so that no obvious r e l a t i o n appeared to exist between elevation and l i c k use. Those l i c k s which are situated near the bunchgrass zone or i n the path of the spring migration have a greater tendency to-ward joint use than those which occur i n heavy timber. This leads to the Idea that the vegetative zone supplying the best winter range increases joint use. Joint use occurred only i n the spring when ungulates were concentrated on areas of new spring growth. As sheep, deer and elk moved to the i r summer ranges joint use decreased i n i n t e n s i t y . 3k INFLUENCE OF SEASONAL PATTERNS ON EXTERNAL PARASITE LOAD, PREPATION AND HUNTER KILL During the spring when big game species are centred i n the bunchgrass zone and Douglas f i r parkland, the paralysis t i c k comes out of hibernation. It attacks game animals i n the spring of the year from about March to May, with a few s t i l l active i n June, The goat i s present i n t h i s vegetative zone at t h i s time. Males trapped i n early spring bore large num-bers on most of their body; approximately six i n an area covered by the palm of one hand. It has not been shown that th i s t i c k causes Rocky Mountain spotted fever i n big game but they may be of serious consequence, by draining the host's v i t a l i t y when i t i s i n an already weakened condition from winter food shortages (Cowan, 195Do Predation Cowan and Brink (19^-9) state that an important re-s u l t of the a t t r a c t i o n that l i c k s exert upon game i s the ex-posure to predation that ensues. Almost a l l goat k i l l s that they found were adjacent to l i c k s . During two summers f i e l d work at six l i c k s no goat k i l l s or remains were found. The three low e l e v a t i o n a l l i c k s (Lazy Lake, Toby Creek, Stoddart Creek) have such predators as black bear and cougar. The high e l e v a t i o n a l l i c k s have these two predators as well as g r i z z l y bears. Black bear were 35 seen at the low and high e l e v a t i o n a l l i c k s ; g r i z z l i e s at some high e l e v a t i o n a l l i c k s and cougars were not seen at either, however fresh cougar tracks were seen near the Lazy Lake l i c k . Although many t r i p s were made through regions 1 and 2 and the l i c k , at the various l i c k s i t e s throughout the summer, no evidence of predation was found e On numerous occasions black bear were seen at a l l l i c k s . The female goat trapped during late July at Toby Creek was kept i n a pen for two weeks near the l i c k . On three separate occasions black bear were seen feeding on Shepherdia berries (buffalo berries) within 100 feet of the pen. The goat did not seem disturbed by the bear's presence and the bear did not bother the goat. These were not repeated observations on the same bear. The cougar population has been reduced i n th i s region due to bounties and paid predatory hunters; causing a decrease i n cougar predation. It seems s i g n i f i c a n t that only one set of cougar tracks was found during two summers of f i e l d work 8 Predation does not seem to be a very important neg-ative factor a f f e c t i n g l i c k use. E f f e c t of the hunting season on l i c k use The termination of l i c k use was rel a t e d to the opening day of hunting season for the two summer study periods,, In 1965s there were reports of approximately 8 to 36 10 goats shot i n l i c k s or i n areas adjacent to them,, Although this i s not extremely high, i t i s important because most animals were females, A female was shot at Lazy Lake on September 8, one at FIndlay Creek on September 1, three at Dutch Creek and two females and a yearling i n the v i c i n i t y of the E l k Creek l i c k . The opening day of hunting season was August 2M-. These l i c k s make up a very small part of the t o t a l number present i n the East Kootenay, In 1966, the hunting season did not open u n t i l September 3» No reports of goats k i l l e d i n the l i c k s were received by September 20, although many people said that they had hunted the l i c k s . It seems that l i c k use had been t e r -minated and the goats had moved to other ranges. The l i c k s at a low elevation had a lower goat k i l l during both summers than the l i c k s at a high elevation. The later opening of the hunting season shows a reduced k i l l i n the l i c k s although temperature and weather may a f f e c t this a l s o . 37 PATTERNS OF DIFFERENTIAL USE IN TIME I have shown that both sexes of the mountain goat use l i c k s during the summer. It i s the purpose of this section to explain d i f f e r e n t i a l use w i t h i n - l i c k s and between l i c k s . The four month seasonal use pattern can be divided into shorter periods when It i s r e a l i z e d that each sex has Its own pattern of use. This i s termed d i f f e r e n t i a l use i n time and implies that c e r t a i n factors are l i m i t i n g male use to a speci-f i c time and female use to a s p e c i f i c time, within a l i c k . The d a i l y t o t a l of males and females at the Toby Creek l i c k was plotted through May, June and July as shown i n Figures 1U-, 15 and 16. The great fluctuations i n numbers from day to day were caused by animals moving into and out of the l i c k synchronously with weather changes. Licks at lower elevations were used by males i n late A p r i l and early May while those at higher elevations were not used u n t i l late May, During i n i t i a l use, numbers were small but increased over the next 10 to 20 days. Prom Figures 1*+, 15 and 16 showing d a i l y numbers, cumulative t o t a l s were plotted for 10 day in t e r v a l s from May to August, as shown In Figure h0 The peak for male use occurred around the end of May and for female use around the end of June or f i r s t part of July. The overlap of use by the sexes between May 30 and June 30 indicates the change-over Males and yearlings Females, kids and yearlings 1 0 0 9 5 9 0 8 5 8 0 7 5 7 0 6 5 6 0 5 5 -1 < 5 0 4 5 z < 4 0 OF 3 5 OS 3 0 111 ca 2 5 s 2 0 3 Z 15 10 5 TO" 2 0 May 3 0 10 2 0 June 3 0 10 2 0 July 3 0 1 0 August TIME IN TEN DAY INTERVALS FIGURE 4. The cumulative totals for 10 day periods,at the Toby Creek lick, showing differential use in time. 38 from male to female use. The change occurred g r a d u a l l y ; num-bers of both sexes were reduced and the m a j o r i t y of females e n t e r i n g the l i c k had k i d s w i t h them. The cumulative t o t a l s of males and females are sim-i l a r but a f t e r the peak, female numbers d i d not drop as f a s t as male numbers. The peaks reach approximately the same height and the buildup i n both sexes took about 20 days, Male use occurred between May 1 and J u l y 10, a period of 71 days and female use l a s t e d f o r about 70 days. T o t a l use was approximately equal f o r males and females but i n t e n s i t y of use by each sex as a whole could not be measured. I n d i v i d u a l males and females could not be compared as to the degree of a c t u a l l i c k i n g t a k i n g p l a c e . I f the animals are attempting to s a t i s f y a p h y s i o l o g i c a l d e f i c i e n c y then i t would seem that the same time at the l i o k serves to s a t i s f y eaoh sex, r e g a r d l e s s of d i f f e r e n c e s i n the p h y s i o l o g i o a l s t a t e . Comparing F i g u r e s 5 and 6 shows the r e l a t i o n of k i d d i n g t o changeover of use by sex. As female numbers i n -crease i n June §o do k i d numbers. She f l u c t u a t i o n s i n female numbera i n F i g u r e $ c l o s e l y p a r a l l e l those i n k i d numbers shown i n F i g u r e 6. This suggests that females were r e s t r i c t e d from using the l i o k due to k i d d i n g . Very few (2 or 3) females without young were i n the l i c k p r i o r to the i n f l u x of females wi t h young. This appears reasonable, since females without 20 1 1 2 3 4 5 6 7 8 9 lO 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 June TIME IN DAYS FIGURE 5 . The increase in female use of licks during June, after the k idd ing per iod. 9 tt U 8 tt Ul nr 111 ft! U >- 6 CO O X 5 < 4 O ui CO 2 29 30 31 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 May June TIME IN DAYS FIGURE 6. The increase in number of kids at the Toby Creek lick in J u n e . 39 young were associated with females with young as an i n t e g r a l part of the family group and very seldom t r a v e l l e d alone. Be-havioural factors rather than kidding would appear to r e s t r i c t these animals from using the l i c k . Females with and without young moved into the l i c k i n groups after kidding. Brandborg (1955) suggests that there i s a rapid increase In the average group size as the kidding period progresses. It may be that d i f f e r e n t i a l use serves to reduce antagonism between the sexes during kidding. Brandborg (1955) suggests that no tendency was observed"for the nannies with kids to j o i n b i l l i e s i n the formation of close family groups after the kidding period. At t h i s time the female becomes aggressive toward the male and protects her young. D i f f e r -e n t i a l use may be an e f f e c t i v e means of separating the aggressive female from the non-aggressive male. D i f f e r e n t i a l use as a general trend It has been shown that d i f f e r e n t i a l use i n time, as It occurs at the Toby Creek l i c k i s a w i t h i n - l i c k event, not to be confused with separation of the sexes between l i c k s , which i s to be explained. The separation of the sexes at this^ l i c k i s d e f i n i t e . As a trend that can be applied to other l i c k s , i t appears that distance and the complexity of the l i c k , i n terms of habitat features, have a modifying influence. The data shown i n Figures 7, 8 and 9 suggest that 30 28| 26 «A 24 ^ 22 Z z < u . o cc u i ca * 20| 18 16 14 12 10 8 6 4 2 Males Females Apri I May June July TIME IN M O N T H S August FIGURE 7. Differential use in t ime at the Dutch Creek l i ck , M a l e s Females < Z < o tt Ul 30 28 26 24 22 20 18 16 14 12 10 8 6 4 2 Apri l Mai June July TIME IN MONTHS August FIGURE 8. Differential use in t i m e a t the Stoddart Creek l i ck . Males Females 24 22 </) 20 S 16 Z 14 u. 1 2 O io ui 8 00 i 6 i « 2 Apri l May June Ju l y Augus t TIME IN M O N T H S FIGURE 9. Differential use in t i m e at the Lazy Lake lick. 4-0 d i f f e r e n t i a l use i n time occurs at the Dutch Creek, Stoddart Creek and Lazy Lake l i c k s . The overlap of use by the two sexes occurs i n June; with males observed i n the spring and mainly females i n July and August. As w i l l be shown, the short distances t r a v e l l e d at the Lazy Lake l i c k tend to promote d a i l y t r i p s so that male use extends Into the summer and somewhat obscures the r e s u l t s . In general, i t may be assumed that d i f f e r e n t i a l use probably i s present to some degree at a l l l i c k s i n the study area. Of the four l i c k s having s u f f i c i e n t quantitative data to show d i f f e r e n t i a l use, three are at low elevations and one at a higher elevation. Use of the high elevation l i c k (Dutch Creek) begins later i n the spring and ends la t e r i n the f a l l . The peaks of male and female use have sh i f t e d to later i n the summer with use by females s t i l l occurring near the end of August, Use of the low elevation l i c k s begins e a r l i e r i n the spring and ends e a r l i e r i n the summer. The pattern of d i f f e r e n t i a l use i s similar at each l i c k yet the peaks can s h i f t depending on the elevation. D i f f e r e n t i a l use between l i c k s The separation of the sexes i n time, within a l i c k , has been shown but the p o s s i b i l i t y exists that the sexes could be separated by using d i f f e r e n t l i c k s , During the summer of 1965» young were not seen i n hi the Mary Creek l i c k although males and yearlings were Identi-f i e d . Observations from the summer of 1966 showed females with young to be using the l i c k . None of the l i c k s examined i n the study area appeared to serve one sex exc l u s i v e l y . D i f f e r e n t i a l use i n time as a wi t h i n - l i c k s phenomenon i s present at the l i c k s i n the study area but as a between l i c k s phenomenon, d i f f e r e n t i a l use appears to be nonexistent. The d i f f e r e n t i a l e f f e c t of parasites, joint use, predators and hunter k i l l Since d i f f e r e n t i a l use may be considered a general trend i n l i c k use, i t i s apparent that the above factors w i l l a ffect the sexes unequally. The male i s susceptible to high external parasite loads and i s affected by joint use since i t descends to the v a l l e y bottoms i n early spring. The female does not enter t h i s region u n t i l other ungulate populations have migrated to higher elevations and the heat of summer has reduced the t i c k populations. Both sexes come into contact with a wider var i e t y of predators present i n this region. I t i s possible that the greater va r i e t y and number of predators could increase ths mortality rate. Cowan and Brink (19^9) r epo r t tha t almost a l l goat k i l l s hav@ been found adjacent t© l i e k § 9 In tho study area predator c e n t r e ! has probably 2"@«* dueed the importance of t h i s f a c t o r . Females w i t h young tend to remain at the l i c k s l a t e r i n the f a l l , making them more vulnerable to hunters. The degree to which each of these factors reduces the population i n t h i s area i s not known. ^3 G R O U P SIZE IN THE LICK AS INFLUENCED BY DIFFERENTIAL USE Males and females using the l i c k at d i f f e r e n t times during the summer showed a pattern of grouping that was con-sis t e n t with t h e i r behaviour. The fact that males and females are very r a r e l y seen together except during the r u t t i n g period and that yearlings are spurned by th e i r mothers i n the spring are t r a i t s influencing patterns of grouping. Although l i c k s served to concentrate aggregations of animals, groups remained small with male groups smaller than female groups, E t k i n (196*+), d i f f e r e n t i a t e s between groups and aggregations by saying that groups are s o c i a l because the members stay together as a r e s u l t of th e i r responses to one another rather than by responses to the other factors i n th e i r environment. He suggests that where i t i s impossible to decide because of lack of evidence, whether a group i s t r u l y s o c i a l or not, the noncommittal term "group'* may be used. In t h i s study, group size was calculated from those animals seen i n the l i c k and those moving between region 2 and the l i c k . An attempt was made to ascertain s o c i a l groups but with males the noncommittal term "group" i s probably more appropriate. Female family groups were usually considered s o c i a l since they remained a basic u n i t . The female family group contains a female and a k i d , usually one yearling and often a female without a k i d . Since we are considering groups rather than aggregations i t seems appropriate to include kids i n the c a l c u l a t i o n of female groups, i n order to show a d i f f -erence between male and female groups. Yearlings were also included i n the c a l c u l a t i o n of group size although i t was almost impossible to d i f f e r e n t i a t e sex. In only one instance were yearlings sexed. Two yearlings entered the l i c k with a group of 3 females and 3 kids and one female without a k i d . One yearling was male and one female, It i s not to be con-cluded that when females with newborn young chase yearlings away, that male yearlings enter male groups and female yearlings enter female family groups. Observations showed that the number of yearlings, regardless of sex, were approximately equally divided between adult male and female groups. In defining a group as such, we hope to show that we are r e a l l y dealing with group size rather than numbers and i n t h i s way can d i f f e r e n t i a t e a group from an aggregation of any number of separate animals a l l on the l i c k at once. Group s i z e was calculated as average t o t a l group s i z e per month, as well as eaoh sex determined separately. T o t a l group s i z e The average t o t a l group s i z e i s shown i n Figure 10. I t shows a steady increase from A p r i l to J u l y with the greatest increase between June and J u l y . This increase i n April May June Ju ly August Sep tember TIME IN MONTHS FIGURE 10. The a ve r age tota l group size (all sex and age c a t e g o r i e s ) per month, using observa t ions f rom al l l i c k s . ^5 July probably represents the changeover to female use, with the female family group becoming prevalent. The A p r i l r e s u l t s are based on a limited number of observations but a t o t a l of 2H-3 observations was used to determine average monthly values. The decrease i n average group size between July and September i s probably the r e s u l t of large groups returning to the summer range, leaving i n d i v i d u a l family groups or females with young, Male group size The average group size for males, as shown i n Figure 11, was calculated from 152 observations, Male group size was greatest i n A p r i l and May but decreased from June to July, It appears that when males are concentrated i n region 2, i n the spring, group size i s larger than i n June when the male num-bers are reduced. This would suggest that we have aggregations In the spring due to crowding. Since males and females are present i n region 2 from June to July, crowding would also be present, yet i t i s at this time that male group size i s r e * ducedo Observations showed that males did not attempt to j o i n female family groups t It may be possible that male group sine is a function of the number of males present in an area re-gardless of the prepense of females Brandborg (1900) iuggggti that larger group s i i a s en the Bid Butts 3?angei (7.5 and 6*1 aniiaale p§r group f o r July and August) may b§ attributtd to the g r s a t e r population d s n s l t y en t h i s rangis April May June July August September TIME IN M O N T H S FIGURE 11. The average group size for males (all age categories ) per mon th , using observat ions from all l i cks . h6 Female group size The average female group s i z e , as shown i n Figure 12, was calculated from approximately 91 observations. The largest group size i s i n July and represents intensive use by the female family group» The increase i n group size from June to July i s attributed to the Increase of family groups and units of family groups 0 In May, limited observations may bias t h i s f i g u r e . Female groups f i r s t to enter the l i c k and those la s t to leave were usually single family groups or females with kids, thus in d i c a t i n g a reduced group size at the time,, It i s known that the peak of reproduction i n most ungulate pop-ulations i n t h i s region occurs during a 2 to 3 week period i n the spring. Fewer females have young before and after the peak than during i t . As stated, single family groups were most prevalent during i n i t i a l l i c k use and again when l i c k use terminated. The gradual increase, peak and gradual decrease seen during reproduction i s similar to that of female use of the l i c k s and may serve to explain reduced group size during early and late l i c k use. The evidence i s strong that females that are to have young do not come to the l i c k u n t i l after these are born. The reluctance of the pregnant females to leave t h e i r normal range u n t i l after p a r t u r i t i o n , taken along with the cohesion of female groups no doubt explains the f a i l u r e of even those females that lack young to come to the April May June July August September TIME IN MONTHS FIGURE 12. The average group size for females (all age categories ) per month, using observations from all licks. 5+7 l i c k s e a r l i e r . The t o t a l average group size at the l i c k during June, July and August i s comparable to that observed by Brandborg on alpine ranges, as shown i n Table 1 0 The female average group size more c l o s e l y approximates his data. The reduction i n group size during May and August may be the r e s u l t of a reduction i n density and may not c l e a r l y represent the true group s i z e . It appears that l i c k use does not appreciably a l t e r the group size as the animals move between the l i c k and th e i r summer range. In only two Instances were males seen c l o s e l y associated with female groups i n the l i c k s . It Is thought that they were not part of the group but represented an aggregation, since the males separated from the group at the s l i g h t e s t disturbance. As stated e a r l i e r , Brandborg (1955) suggests that there i s no tendency for the nannies with kids to j o i n b i l l i e s , as suggested by Andersen ( ^ O ) . The occasional occurrence of males i n mixed groups during the s p r i n g and summer indicates that they do not l e a d an e n t i r e l y s o l i t a r y e x i s t e n c e . Th© aggregations of nalee^ e b i t r v i d fending i n r e g i o n were h i g h l y unstable. S e v e r a l i&st&noes wgr© ob-s e r v e d where four or f i v e males were f e e d i n g or moving about on the same c l i f f . The aggregation did not exhibit any s o c i a l t i e s among animals. Other males could enter t h i s s i t u a t i o n and some i n the aggregation leave, causing a complete change. Yearlings were observed i n female family groups and with males i n the spring. It i s believed that females abandon yearlings to have young of the year. Observations during the summer of I966 showed that females with young are antagonistic toward yearlings and i n many cases force inde-pendence upon them. Throughout the summer, yearlings are chased by the females whenever they wander close to a k i d , however, the i r drive to associate with their parent i s s u f f i c i e n t l y strong that they frequently follow along behind the nanny and k i d . Yearlings are also chased by other adults since they are subordinate. About half the yearlings seen were with females with young, showing that the female i s not always successful i n chasing them away. The s h i f t of the yearling from female to male groups and the factors a f f e c t i n g I t , are not known. Yearlings with adult males tend to i n -crease the average male group s i z e . Table 1 A monthly comparison of average group s i z e , using data from Brandborg (1955) and data from the East Kootenay region.  Month Range Average Group Size Range T o t a l Group Size Female Group Size May Salmon River 2.7 - Toby Creek 1.6 2 Red Butte 3.5 - 2.9 B i t t e r r o o t River 2.6 -June Salmon River 3.6 - • Toby Creek 2.3 3.2 Red Butte 3.3 - 3oh S e l k i r k 3.h -July S e l k i r k 3.6 Toby Creek 3.7 h Red Butte 7.5 - 5.2 Selway 3.8 River -Teton River 6,1 -August Salmon River 3.0 - Toby Creek 2.3 2.3 Red Butte 6,1 - 3.9 Selway 2.8 River 50 BEHAVIOUR During the two summers of f i e l d work c e r t a i n aspects of goat behaviour, regarding the use of l i c k s , was thought to be important. In region 2, aggregations of males were continually-changing i n makeup as new aggregations entered the region and others l e f t . When two aggregations met, goats moved from one to the other f r e e l y but at no time was antagonism observed. When males were i n the l i c k , adults were not observed to d i s -place one another from preferred l i c k i n g s i t e s , although year-lings were constantly chased from these s i t e s . Males and females used the l i c k s at d i f f e r e n t times during the summer, however one male - female encounter was observed at the Lazy Lake l i c k . At this time the female chased the male off the t r a i l and made i t detour widely. The male was much larger than the female but did -not r e t a l i a t e when the female charged. Female encounters were common i n region 2 and most l i c k s In the study area. On June 7? 9 goats were observed In the Dutch Creek l i c k . There were 3 females with kids, 1 female without a kid and 2 yearlings. The majority of l i c k i n g a c t i v i t y occurred on the high l i c k i n g s i t e s with females with kids displacing females without kids and yearlings. Females without kids could displace yearlings. Among the females with 51 kids i t appeared that size played a major role i n determining which female was dominant. Although size was determined by observation only, the supposedly larger females could always displace a smaller one from a preferred l i c k s i t e . Aggressive a c t i v i t y consisted of the dominant female making one or two hops, with head lowered, toward the smaller female. The aggressor would bring i t s head up i n a horn thrust on the l a s t hop but contact was not observed. Within the female family group, the same order of dominance prevailed: females with young, females without young and yearlings. This type of hierarchy would benefit females with young by allowing them preferred s i t e s i n the l i c k . In region 2, females often met on the main t r a i l as they moved to and from the l i c k . No evident r e l a t i o n appeared to e x i s t between those occupying a s i t e on the t r a i l and those showing dominance. Animals lying on the t r a i l displaced approaching animals as often as they were displaced. The size of a group approaching or occupying a s i t e did not seem to matter as far as dominance was concerned. In one case, a female and k i d approached a group of 5 goats lying on a t r a i l and chased them off so they could get by. The group consisted of a female with a k i d , a female without a k i d and 2 yearlings. Again i t appeared that size of the i n d i v i d u a l animal was the determining f a c t o r . 52 DIFFERENTIAL USE AND ITS EFFECTS ON MOLT The patterns of seasonal and d i f f e r e n t i a l use of l i c k s as a part of the ecology of the goat show a d e f i n i t e e f f e c t on the rate of molt. This i s evident when we note the termination of molt i n males, females without young and females with young. The habitat conditions and the time each class of animals spends i n these conditions greatly affects the loss of h a i r . Cowan (19^0) describing the molt i n sheep, reports that, "Individuals inhabiting areas where there i s some brushy cover tend to molt the old hair on the sides and abdomen early as a r e s u l t of the rubbing of these parts on the bushes". Cowan (op. c i t . ) further states that, "As i n many, i f not a l l ungulates, physical condition undoubtedly affects time of molt. Mature rams on the average molt e a r l i e r than ewes with young or animals of either sex i n poor condition." It i s not known whether the phy s i o l o g i c a l mechanisms involved i n molt induce molt at the same time i n each class of animal but the following discussion describes the influence of the l i c k on the pattern and termination of molt. By the end of May, males had lost patches of hair on their nose, throat and chest region including the top inside part of their front legs. Hair had also been lo s t from the rest of the body, causing a thinning e f f e c t but no patchiness was evident i n the posterior region. The phys i o l o g i c a l process 53 of molt had begun i n the males but the r a p i d i t y with which they lose their hair r e s u l t s from the abrasion of the pelage by the shrubby t e r r a i n i n which they are l i v i n g . Towards the end of June, males had lo s t approximately one-half of the old coat and the remaining portion was- ex-tremely t h i n and ragged. The molt moves from anterior to posterior. The front quarters and lower portion of the ab-domen are completely bare at thi s time while the hind quarters and upper portion of the abdomen s t i l l r e t a i n the old coat. The females without young have lost approximately 25 percent of their old coat toward the end of June. This i s i n the anterior region, with the sequence the same as for males. Females had just begun to use the l i c k and abrasion by the vegetation had continued for a shorter time. In July, the males and females without young lose the remaining portion of t h e i r old coatj with males losing theirs f i r s t . Both had completed molt by the 15th to 20th of July. At the end of July t h e i r new coat was £ to 1 inch i n length. Brandborg (1955) observing goats on alpine ranges states, "of 10 animals observed at close range on July 1*+ near Red Butte, only one adult male had shed the old hair com-pl e t e l y . " It appears then that abrasion may hasten molt. The females with young lose their old coat much more slowly than males or females without young. The females with 5if young have spent the same length of time i n the v i c i n i t y of the l i c k as females without young but have retained much more of their old coat. Most animals have completed molt by the end of August. The pattern of molt for these three categories i s shown i n Figure 13. Although abrasion aids i n hair re-moval, the o v e r - a l l pattern i s fixed by p h y s i o l o g i c a l processes. S i m i l a r l y , d i f f e r e n t i a l use of l i c k s , i n time, leads to use of dust baths by males and not by females. Males a r r i v i n g at the l i c k i n early spring are molting and become mere susceptible to attack by the paralysis t i c k , Th© animal© would s i t on their haunches and kick d i r t onto their body with their front feet, Gelst (196*+) describes t h i s action i n the adult male during r u t , by sayings "Adult males were soon e a s i l y distinguished from females by black patches of d i r t on the rump, as w e l l as by streaked flanks, matted "trousers" and d i r t y b e l l i e s . These they received from pawing r u t t i n g p i t s , a behavior associated mainly with aggression, but which could appear i n courtship or spontaneously." It seems quite possible that t h i s i s the same be-havioural stance which i s being used to achieve another end. The aggressive tendencies shown by the head and body position are absent. It was thought that males may dig these pits i n order to l i e on the cooler earth. Since females were i n the l i c k during the hottest part of the summer and did not use the 100 9 5 9 0 8 5 8 0 ED 7 5 *— OL 7 0 s 6 5 AT 6 0 0 5 5 (J Q 5 0 O 4 5 o 4 0 •NT 3 5 w 3 0 UJ Q. 2 5 2 0 15 10 5 Males Females without young Females with young April May June July TIME IN MONTHS FIGURE 13. The sequence of molt, caused by abrasion from vegetation, while the animals are using the lick. Auqust September 55 dust baths, th i s did not seem the l o g i c a l answer. Females entered the l i c k after the t i c k i n f e s t a t i o n and so dust baths were thought to be used by the males, to reduce i r r i t a t i o n caused by t i c k s . 56 LICK USE OVER A 2k HOUR PERIOD Intensive lick, use occurred at d i f f e r e n t times of the day at d i f f e r e n t l i c k s , depending on the structure of the l i c k . A comparison of the Lazy Lake, Dutch Creek and Toby Creek l i c k s serves to i l l u s t r a t e the differences. As shown i n Ta„ble 2 intensive l i c k use at both the Dutch Creek and Lazy Lake l i c k s occurred i n the afternoon. Table 2 P e r i o d i c i t y of d a i l y use at the Lazy Lake and Dutch Creek l i c k s , based on observations of groups. No, of, Morning Afternoon Evening Lick Groups 0/0 0/0 0/0 Lazy Lake 30 0 83 17 Dutch Creek 21 0 67 33 The goats did not use the l i c k i n the morning but some use occurred i n the evening. These two l i c k s have been described as simple l i c k s having few habitat features common to region 1« This r e s u l t s i n a p e r i o d i c i t y of use not seen i n more complex l i c k s . The fact that more l i c k i n g was done In the evening at the Dutch Creek l i c k can probably be attributed to the short distance between region 2 and the l i c k . At Dutch Creek, large groups (9-15) used the l i c k mainly i n the afternoon with i n -d i v i d u a l animals entering i n the evening. At Toby Creek, In-tensive l i c k use showed less r e s t r i c t i o n to the afternoon. 57 Animals were observed l i c k i n g i n the morning, evening and afternoon, although the majority of use s t i l l occurred i n the afternoon. This i s considered the most complex l i c k and animals would spend from 2 to h days i n the l i c k at one time. Since the animals could feed and move about the l i c k from dawn t i l l dark, l i c k use was not r e s t r i c t e d to any one period of the day. The number of habitat features, then, plays an important role i n concentrating l i c k use during the daylight hours. 58 SEQUENCE OF LICK USE DURING THE SUMMER It has previously been shown that males entering region 2 i n A p r i l and May u t i l i z e succulent forage before entering the l i c k . As the summer progresses, animals feed at successively higher elevations so that they are continually feeding on young forage while using the l i c k . As shown i n Table 3 animals i n the v i c i n i t y of the l i c k were categorized as to whether or not they were i n the l i c k . During A p r i l , males were not observed i n the l i c k and almost a l l observations showed them to be feeding. As the summer progressed and the distance between succulent forage and the l i c k increased, animals were more frequently observed l i c k i n g once they reached the v i c i n i t y of the l i c k . Males, i n the v i c i n i t y of the l i c k i n spring, were there for two reasons; to feed on spring forage and to use the l i c k . Males were ob-served using the l i c k more frequently as the time spent feeding on succulent forage increased. Females, feeding i n region 1 and the higher portions of region 2, upon entering the v i c i n i t y of the l i c k i n June and July were more often observed i n the l i c k as th i s was their primary reason for being at t h i s low elevation. It appears that as the cumulative time animals had been feeding on succulent forage increases and as the distance between succulent forage and the l i c k increases, l i c k use i n -creases. This i s shown by the observation that i n June and 59 July animals were more frequently seen i n the l i c k than merely i n the v i c i n i t y (Table 3). Upon examining Tables 2 and 3 i t i s seen that l i c k use was greatest i n the afternoon and most intensive i n July. It was during July of both summers that the temperature was highest and number of storms reduced. These statements w i l l be examined i n more d e t a i l i n a later section. As l i c k use increased over the summer i t must be re-membered that group size also increased. Increase i n l i c k use may be a function of the larger groups being more v i s i b l e . Table 3 Change i n degree of l i c k use, using combined group observations from the Dutch Creek and Lazy Lake l i c k s . Month No. of Groups In the Lick 0/0 Not i n the Lick 0/0 A p r i l i+ 0 100 May 8 12.5 87.5 June 21 28 72 July 20 80 20 60 DISTANCE AS IT RELATES TO LENGTH OF TIME SPENT IN THE LICK Distance seems to be important i n determining the amount of time spent i n the l i c k at any one v i s i t . This can best be shown by comparing the Lazy Lake l i c k to the Dutch Creek l i c k . This i s , i n essence, a comparison of a l i c k where a short distance separates region 1 from the l i c k to one where a much longer distance separates region 1 from the l i c k . At Lazy Lake, observations show that reduced amounts of time were spent i n the l i c k regardless of the time of day l i c k i n g occurred. Most observations were of female groups and the average time was 2 hours. It appears that the a c c e s s i b i l i t y of the l i c k has reduced the amount of time spent i n the l i c k although i t i s l i k e l y that frequent t r i p s may o f f s e t t h i s . At Dutch Creek, the average amount of time spent i n the l i c k i s about 5 hours as shown i n Table h. Table h The average time spent i n the Lazy Lake and Dutch Creek l i c k s based on observations of female groups. No. of Average Time Lick Observations In the Lick (hr.) Lazy Lake 9 2 Dutch Creek 13 5 Again most observations are of female groups so the comparison i s v a l i d . In the spring, males were moving between region 2 and the l i c k and spent very short periods i n the l i c k , much the 61 same as at the Lazy Lake l i c k . In general^ i t seems that an-imals which t r a v e l long distances to reach a l i c k t r a v e l i n larger groups, are regular i n their timing and spend longer periods i n the l i c k at any one time. The reverse i s true of situations where short distances p r e v a i l between region 1 and the l i c k . Complexity of the l i c k Although distance appears to be important i n reg-ulating group s i z e , time of entry to the l i c k and amount o f time spent i n the l i c k , we must consider complexity of the l i c k as a regulating factor once the animals reach the l i c k . At Toby Creek, animals spend 2 to h days In the l i c k although the distance between region 2 and the l i c k i s very short. While i n t h i s l i c k goats feed and bed much as they do on the i r summer range so that less time Is spent ac t u a l l y licking„ It appears that once i n the l i c k , l i c k i n g , feeding and bedding are of equal importance« The Butch Creek l i c k has few habitat types and once the animals are i n the l i c k , l i c k i n g i s of primary importance« Apart from occasional drinks and the odd departure from the l i c k to urinate and feed, most time i s spent l i c k i n g * The Lazy Lake l i c k has l i t t l e feed i n the l i c k and although some escape t e r r a i n i s present goaty did not appear to use i t for bedding purposes 6 When goats were i n this l i c k the majority of time was spent l i c k i n g . 62 In conelusion 9 i t may be said that although the de f i n i t e use of t r a i l s increases the e f f i c i e n c y of movement between regions, distance, and complexity of the l i c k regulate the movement. 63 PATTERNS OF FREQUENCY OF USE It i s believed that the frequency of use by an i n -d i v i d u a l or group i s largely determined by the nature of the factors i n i t i a t i n g l i c k use, It has been postulated that succulent vegetation (Dalke eJ^aJL, 1965) and- or a dietary d e f i c i e n c y ? (Cowan and Brink, 19*+9 and Honess and Frost, 19*+7) could lead to i n i t i a t i o n of l i c k use. This suggests that i f i t Is due s o l e l y to a winter dietary deficiency, the animals should enter the l i c k , s a t i s f y the craving and leave. If i t i s due to a winter dietary deficiency plus a high water uptake on a low s a l t diet i n the spring, the animals w i l l probably make frequent t r i p s to the l i c k . Since animals feed on succulent vegetation prior to using the l i c k , the effects of a winter dietary deficiency could not be determined. The i n i t i a t i o n of l i c k use could only be r e l a t e d to the c r i t i c a l period i n the spring. In order to ascertain frequency and duration of use a limited program of marking was undertaken at the Stoddart and Toby Creek l i c k s . This -was set up to allow us to describe patterns of movement, to assess the r e l a t i v e Importance of region 2 , to relate animal movement to temperature and storms, to deduce changes i n the population and eventually give a population estimate, A t o t a l of 2 0 animals was marked at the two l i c k s 6h and subsequently i d e n t i f i e d . Others were known to have been marked by our automatic marking devices but we were unable to confirm t h i s by actual sightings. Marking was car r i e d out i n the l i c k for short periods during intensive l i c k use and was continued throughout the summer. The r e s u l t s of marking and subsequent sightings are shown i n Table 5o Duration Within each major period of use by the sexes, an attempt was made to determine the duration of use by indiv -i d u a l s . This would indicate whether populations using the l i c k changed i n number and composition 0 The duration an i n -d i v i d u a l spent i n region 2 or 3 could not always be measured accurately, since the period animals were present prior to being marked could not always be determined. The time between the known marking and subsequent sighting i s taken as the minimum time the animal was present i n region 2 or 3, Inten-sive marking was accomplished during the early part of May, the l a t t e r part of June, and the l a t t e r part of July, The periods between marking were used to note changes i n the pop-ulation,, As shown i n Table 5? animals marked i n the l i c k i n early May were present from 2 to 7 days In the l i c k or region 2b One animal marked on May 12 was present for 16 days but was not seen again after May 27* During the l a t t e r part'of Table 5 Marked animals and subsequent sightings at the Stoddart and.Toby Creek licks„ Subsequent Sightings Period between Date Marked Seen away Marking and Animal In l i c k Lick Seen i n l i c k from l i c k Sighting l c Adult male May 12 T May 1 8 7 2 0 Adult male May 17 T May 1 8 2 ?° Yearling May 18 S May 19 2 h„ Adult male May 1 2 T May 18-19-20-2^-27 16 5 o Adult May 17 S May 2h & 2 7 May 22 6 6 . Adult male May 27 T 6 7 o Adult male May 23 T May 25 3 8. Yearling male May 26 S May 26 ^ - L . 9 c Adult male Between May 23-27 T May 28 May 27=29 6 10. Adult male Between May 23-27 T May 28 May 27-29 6 11. Adult male May 31 T May 31 1 12. Yearling May 2h S May 27 h 13. Adult male June 13 S June 13 1 Ih. Adult female June 19 T June 29 & 30 12 15 c Adult female June 25 T June 28 if 1 6 . Adult female June 25-27 T June 29 if 17 o Adult female June 2h T June 28-29 June 25-27-28 6 1 8 . Adult female June 2h. T June 25 2 19 c Adult female June 27 T June 29 3 20. Adult female June 25 T June 30 6 T - Toby Creek l i c k S - Stoddart Creek l i c k ON 66 May, marked animals were present for periods of 1 to 6 days. During periods between marking and subsequent sightings, an-imals were thought to spend the majority of time i n the timber and on the c l i f f s i n region 2„ Animals were more consistently observed i n region 2, and on two occasions animals were ob-served i n the l i c k on one pa r t i c u l a r day and In region 2 the days preceding and following it„ In May, males spent much time feeding i n region 2 and appeared to make frequent t r i p s to the l i c k (tracks and shed hair) although sightings were not made to confirm t h i s 0 During the l a t t e r part of June animals were present for 2 to 12 days as shown by marking and sub-sequent sightingso Marked females appeared to t r a v e l between region 2 and the l i c k less frequently than males and spent longer periods i n the l i c k on any one v i s i t than did males (tracks and shed h a i r ) . Frequency and patterns of movement The patterns and frequency of movement between regions 2 and 3 were extremely d i f f i c u l t to determine. Inad-equate sightings prevented us from following i n d e t a i l the movements of very many i n d i v i d u a l animals. Animal number marked i n the l i c k on May 12 was observed feeding i n region 2 for three days (May 18, 19, 20)„ A storm between May 20 and 23 probably did not permit t h i s animal to enter the l i c k but between May 2k and 27 i t was thought to have entered the l i c k . 67 Adult male number 6 was marked on May 22 ( i n the l i c k ) and was again observed i n the l i c k on May 2*+ and 2 7 , It was not ob-served i n region 2 between these dates. Two males (numbers 9 and 10) marked i n the l i c k around May 2*+ were sighted i n region 2 on May 2 7 , i n the l i c k on May 28 and again i n region 2 on May 2 9 , They were together at a l l sightings. An adult female, marked on June 19 was observed i n the l i c k on June 29 and 3 0 . Between these dates i t s whereabouts were not known exactly but i t was thought to be In region 1 or 2 , Female number 1 7 , marked i n the l i c k on June 2*+, was seen i n region 2 o n J u n e 2 5 5 27 and 28 and i n the l i c k on June 28 and 2 9 . It appears from these data that animals move between region 2 and 3 repeatedly during the period they are i n the v i c i n i t y of the l i c k . They seem to spend considerable time i n region 2 feeding, but make frequent t r i p s to the l i c k . Animals were observed to frequent the l i c k on several occasions during short periods but d e f i n i t e patterns could not be established due to a lack of consistent sightings of marked animals. The importance of region 2 The use of region 2 as a base from which goats move to use the l i c k s was not r e a l i z e d u n t i l marked animals were consistently observed returning to this region. E a r l i e r studies do not mention such an idea and i t was at f i r s t be-lieved that animals were passing d i r e c t l y between the alpine 68 summer ranges and the l i c k , with short pauses along the way. The subsequent sightings of marked animals i s shown in Table 5» Males, i n May, spent a considerable amount of time feeding i n region 2 and while few sightings 'of these animals were made i n the l i c k , tracks and shed hair on the t r a i l leading to the l i c k attested to their frequent movement between the two regions. At i n t e r v a l s along the t r a i l leading to the l i c k , tracks were brushed off each day and hair removed from branches. Certain branches were positioned to remove molting hair from animals entering or leaving the l i c k 0 In many cases, a marked animal which disappeared from region 2 was thought to be i n the l i c k since marked hair was picked up on the t r a i l , Indicating that i t had passed th i s spot and was headed for the l i c k o Males were not observed feeding around the l i c k and returned to region 2 to bed and dig dust baths. It appeared that region 2 was supplying males with a l l their habitat essentials and thus detracted from time spent i n the l i c k o When females entered the v i c i n i t y of the Toby Creek l i c k they fed i n the highest portion of region 2. This i n -creased the distance between the l i c k and area of intensive feeding. Females were more often observed feeding around the l i c k , bedding i n the l i c k and seemed to spend more time i n the l i c k on any one v i s i t . It appeared that they moved less 69 frequently between region 2 and the l i c k , as shown by tracks and shed h a i r . S i m i l a r l y , at Dutch Creek, females made fewer t r i p s between region 1 and the l i c k , did not u t i l i z e region 2 (the vegetation there had matured) and spent considerably longer periods i n the l i c k at any one time, than did the males. Region 2 appeared extremely important as long as i t contained succulent vegetation and the distance between i t and the l i c k was quite short. Environmental factors a f f e c t i n g movement Although mineral d e f i c i e n c i e s and water content of the vegetation may determine o v e r - a l l patterns of l i c k use, such environmental factors as temperature and weather may determine a more refined d i s t r i b u t i o n of use during the summer. Daily temperature for July, 1965? "was plotted as shown i n Figure 17 and for May, June and July of I966, as shown i n Figures 1*+, 15, and 16, The d a l l y readings were obtained from the Cranbrook Airport s t a t i o n i n most cases but also from the Radium Junction s t a t i o n when available (Dept, of Transport, 1965 and 1966), F i e l d c l i m a t i c stations were not set up i n thi s study as only the trends i n temperature were needed and the readings from the two established stations were adequate. In general, the peaks i n temperature, from 65°F0 to 95°F, were warm and sunny with only the occasional thundershower, The lows i n temperature, from 50° F, to 65°F0 approximately, were 70 usually c o o l , cloudy and often stormy. It rained often during the stormy periods. On the above l i s t e d f i g u r e s , animal num-bers were superimposed to show a c o r r e l a t i o n between tem-perature, weather and animal movement. In July, 1965j animal numbers were recorded at the Dutch Creek l i c k . As shown i n Figure 17, at least one large group of animals used the l i c k during each peak i n temperature. A c t i v i t y was greatest at thi s time and tracks showed that for 2 to 3 days during each peak, animals were moving i n and out of the l i c k . Animals were observed to enter the l i c k during the hottest part of the afternoon and remain there for 6 to 8 hours, suggesting a strong c o r r e l a t i o n of l i c k use and tem-perature. During the lows, although dropping only to 65 F., storms were common and very few goats were observed. A c t i v i t y i n the v i c i n i t y of the l i c k was almost n i l with only the odd i n d i v i d u a l animal moving about. The difference i n a c t i v i t y i n region 2 and 3 prior to a storm and during i t , i s shown i n Table 7. Beeman C1957) s. states that i n the lower Selway River area, r a i n storms may come quickly and la s t for days e E l k a c t i v i t y was noted to drop sharply during these r a i n s • The high degree of c o r r e l a t i o n achieved at the Dutch Creek l i c k i n 1965 d i d not occur a t the Toby Greek l i c k i n 1966'as seen by comparing Figures 1^ , 15 and 16 t o Figure 17, This was due mainly to the gr e a t difference i n iummera between TEMPERATURE IN DEGREES FAHRENHEIT NUMBER OF ANIMALS TEMPERATURE IN DEGREES FAHRENHEIT TEMPERATURE IN DEGREES FAHRENHEIT 10 H » ^ tO » g £ H 5 J K M M - g g jj NUMBER OF ANIMALS TEMPERATURE IN DEGREES FAHRENHEIT NUMBER OF ANIMALS TEMPERATURE IN DEGREES FAHRENHEIT © C 70 i i a « ft o c ft ft 3 ft ft 3 3 1 ' I * z: 3 m I * 5 5 a 2 03 O H I 03 03 at < CO CO Cfl 09 03 £• r CD 00 a 5 a 8 09 J2L 03 O o rf* 01 Ol Ol 0) o in co -a cn O -a cn 00 cn CO CO Q cn cn 09 \ / • \ ^1 \ • s ft o •* •* D _ . •+ 3 C "l C » in ~— 9 2 o X 3 c 3 D 3 3 Q Is O fc - Q _ 3 * 3 o 3 Ui Q 3 C 3 or » •I ft (Q 5' 3 VI 09 O 09 09 09 09 CO 09 *. 09 Ol 09 03 09 -I 09 oo to CO 03 O \ \ \ / 71 the two yearso In general, the summer of I966 was warmer and sunnier, fluctuations i n termperature were less pronounced, during the l a t t e r part at least and storms were less common than during the summer of 1965« In Figure i h we see that use of the Toby Creek l i c k follows the fluctuations i n temperature quite c l o s e l y during the l a t t e r part of May0 During the f i r s t part of May, Intensive l i c k use had not yet begun,, This i s due largely to the lower average temperature, the greater fluct-uations of temperature and the frequent occurrence of storms which correspond to the drops i n temperature 0 The c o r r e l a t i o n i s thrown out of phase i n June as average temperature r i s e s and fluctuations are less evident 0 The changes In animal num-bers follow more c l o s e l y the pattern of storms 9 since tem-perature and storms can occur out of phase as wello This i s shown i n Table 6„ Table 6 The number of animals observed i n region 2 and 3 during a period when a storm and temperature are out of phase„ Prestorm Storm Post storm June June June Kegion 5 7 8 9 10 11 2 3 m tH H ! O «S a 0 d 523 «3 5 0 1 2 9 5 3 0 0 0 11 10 72 Very few animals were observed on June 9 o r 10 during a storm, although temperature remained f a i r l y high. On June 11 when the temperature dropped, animal numbers rose and though the weather was cool and cloudy 9 i t was not stormy. Although there were many peaks i n temperature , during June and July, l i c k use was not s o l e l y r e s t r i c t e d to these times and as stated, the degree of c o r r e l a t i o n became less pronounced as the summer progressed. The highest number and greatest a c t i v i t y was seen after storms and during periods of higher temperature. Prior to a storm, observations showed a decrease i n numbers as shown i n Table 7? ^or the l a t t e r half of Mr.y. After a storm, numbers usually increased for 2 to 3 days. The Dutch Creek and Toby Creek l i c k s d i f f e r i n the Table 7 The movement of animals between regions 2 and 3 at the Toby Creek l i c k as a r e s u l t of a severe storm. Prestorm Storm Post storm May May May Region 18 19 20 21 22 23 2h 25 2? 2 3 ; o"; o m ; ° «H H O 'Q -cd 10 12 5 2 3 3 0 1 0 0 6 1 10 8 9 1 2 8 degree to which temperature and animal fluctuations coincide with storms. At Dutch Creek, animals returned to region 1 during a storm and In July no animals were observed i n the l i c k under stormy conditions. As shown by Tables 6, 7 and 8, 73 animals were not seen i n the Tohy Creek l i c k during a storm but were o c c a s i o n a l l y observed i n r e g i o n 2. Table 7 demon-s t r a t e s the e f f e c t s of a severe storm on animal numbers and Table 8 The movement of animals between r e g i o n 2 and 3 at the Toby Creek l i c k as a r e s u l t of a moderate storm. Prestorm Storm Post storm June June June Region 25 27 Morning of 28 Afternoon of 28 29 30 2 3 ;o rt • E O H 18 5 0 0 0 0 6 9 10 6 10 10 movement. I f storms occurred i n quick s u c c e s s i o n , numbers would f l u c t u a t e r a p i d l y as shown In Fig u r e 15. Table 9 com-pares animal use days p r i o r to a storm and during i t . The d i f f e r e n c e i s qu i t e apparent and s t r i k i n g and shows the extent to which l i c k use i s c u r t a i l e d by storms. During a storm Table 9 The e f f e c t of a storm on animal use days of r e g i o n 2 and 3. P e r i o d Days of Observation Numbers Observed In L i c k Or Region 2 Animal Use Days Prestorm 8 80 10 Storm 5 if Post sfcorm 6 117 1^.6 animal movement ceased almost completely but since animals were observed l n r e g i o n 2 before and a f t e r the storm, i t was 7^ supposed that the majority were present there during the storm. The e f f e c t s of a moderate storm as shown i n Table 8 demonstrate t h i s l a s t statement. On t h i s occasion, a storm occurred i n the. morning with a warm sunny period following i n the a f t e r -noon. Animals observed i n region 2 prior to the storm were observed there after i t . The slow buildup of numbers following a severe storm Is harder to explain. The p o s s i b i l i t y exists that some animals move to more protected areas, of which the location i s not known. After the storm they supposedly return to region 2 or move to region 1, The i n i t i a t i o n of intensive l i c k use In May, at the Toby Creek l i c k was rapid and occurred immediately after a storm. As shown i n Table 7 numbers were quite high from May 18 to May 20. Prior to t h i s , from May 10 to 16, there was a period of cold stormy weather. Preceding t h i s , goat numbers at t h i s l i c k had been extremely low. May 17 was cloudy but warmer and the storm had evidently broken. As explained, there was a large buildup of animals from May 18 to 20. This was p a r t i c u l a r l y impressive since 2 days previous there were very few goats i n region 2. As shown i n Figure 16, animal numbers fluctuated although temperature remained f a i r l y high, constant and storms were infrequent. Over t h i s long period of constant weather, animals were free to move between the 3 regions and i t does 75 not seem unreasonable that these fluctuations should ocdur. The fluctuations indicate that changes i n the number and com-position of the population are occurring„ From these examples, i t was thought that temperature and weather play an important role i n regulating the movement of goats between regions 1, 2 and 3« It appears that inclement weather r e s t r i c t s goats from using the l i c k s and region 2 serves as a place to wait out the storm, Changeovers i n the population and factors a f f e c t i n g i t Marking served as an excellent technique to note changes i n the population, the time, at which these changes took place and the possible factors regulating them. Animals were marked i n early May, as shown i n Table 5» and following t h i s , i n early June, unmarked animals were observed i n regions 2 and 3. Males marked i n May were not observed i n June and i t was supposed that some form of changeover had occurred. It i s not known for sure whether the change was abrupt or whether new animals gradually replaced the e a r l i e r ones at the l i c k . The l a t t e r explanation seems more plausible at t h i s time. As stated e a r l i e r , Intensive use of the Toby Creek l i c k began after the storm of May 16, The storm of May 21 caused a c e r t a i n amount of changeover since some animals marked prior to i t were not observed after i t . It was not u n t i l the stormy period at the end of the month that a complete changeover \ 76 occurred. Since intensive l i c k use began just prior to the storm of May 21 and since animals were not i n the l i c k during the storm, i t appears that most had not spent a s u f f i c i e n t time i n the l i c k and remained u n t i l the end of the month. It appears that the storm of May 31 affected the changeover at thi s time but.more s p e c i f i c a l l y , i t s r e l a t i o n to movements of animals between regions 1, 2 and 3 i s not known. During early June, numbers were quite high but animals were unmarked. Females began entering the l i c k i n late June, causing the male - female changeover as explained i n the section on d i f f e r e n t i a l use i n time. This changeover was attributed to the lack of movement prior to kidding. During the l a t t e r part of June females were marked. In July, these animals were not observed i n the l i c k , i n d i c a t i n g another changeover had occurred. At t h i s time, animals using the l i c k were a l l unmarked. Marking began again near the end of July, as use. of t h i s l i c k terminated for the summer but no resu l t s were obtained from t h i s . The changeover of females. can again be related to a severe storm at the beginning of July. It appeared that t h i s storm moved.one population of females out of the l i c k , while another entered after the storm. Observations showed that animals moved during good weather, between regions 1, 2 and 3 with storms between these periods c u r t a i l i n g a c t i v i t y In the l i c k . 77 Population estimate Marking served as a crude method to estimate the population of animals using the Toby Creek l i c k . It has been suggested that approximately 3 to h changeovers of animals took place during the summer and this served as a basis to estimate the number of animals using the l i c k during the summer. Local reports had suggested that only 15 to 20 animals were using the Toby Creek l i c k each summer. If the highest d a i l y total.from each of the h changeovers i s summed, i t appears that approximately 70 animals were present.at the l i c k during the summer of 1966. Using only one day's t o t a l should insure that these were 70 d i f f e r e n t animals. The winter and summer ranges from which these animals are t r a v e l l i n g i s not known but the distance varies from 5 to 25 miles. I t i s not known i f the animals using the l i c k l a t er i n the summer are those t r a v e l l i n g the longest distances. Marking and f l y i n g to determine ranges did not reveal to which ranges the animals had returned. 78 VEGETATIVE SOURCES OF MINERALS Introduction Animals and plants have c e r t a i n mineral requirements, those of animals being supplied by the plants on which they feed. Thus, the plant removes elements from the s o i l s and makes them available to animals. Plants, can absorb large amounts of c e r t a i n elements out of proportion to their abun-dance i n the s o i l , and animals have a similar but less pro-nounced a b i l i t y to select c e r t a i n elements from the plants. Thus, animals require comparatively large amounts of sodium and traces of cobalt but i t has not been found that these elements are necessary to the l i f e of the plant. The macro-metabolic elements e s s e n t i a l to plant l i f e , include phosphorus, s u l f u r , calcium, magnesium, potassium and i r o n . The micro-metabolic elements Include boron, manganese, copper, zinc, molybdenum and ch l o r i n e . The elements Important to both animal and plant l i f e and most often seen as def i c i e n c i e s , ' a r e phos-phorus, calcium, magnesium, potassium and chlorine with sodium and cobalt often appearing as d e f i c i e n c i e s In animals ( G i l b e r t , 1957)o Carnivorous animals, obtaining s u f f i c i e n t sodium from the f l e s h on which they feed, apparently do not crave t h i s element. Herbivorous animals, lacking a s u f f i c i e n t amount i n their food, may receive i t as a supplement] i n the wild state 79 they may t r a v e l great distances to obtain i t from a spring or l i c k ( G i l b e r t , CJD. c i t . ) . Determination of the mineral element sought by goats from natural earth l i c k s requires the elimination of most macro and micrometabolic elements. Seldom do the plants or animals offer any gross symptoms to a s s i s t i n int e r p r e t a t i o n of a mineral deficiency. In general, where no obvious de-f i c i e n c i e s are expressed as animal pathologies, search for the mineral i n demand from l i c k s o i l s has used two approaches: inference from s o i l analyses and from mineral " c a f e t e r i a s " or sel e c t i o n t r i a l s . There have been no previous studies i n which the use of l i c k s by goats has been the main subject of inquiry. Few authors have attempted to relate chemical com-position of the forage to chemical content of the l i c k , re-gardless of the species of animal involved. It i s probable, however, that data from other wild species using the same vegetation w i l l be I l l u s t r a t i v e i n eliminating c e r t a i n elements as a t t r a c t i n g and serve as support for the a t t r a c t i n g element. Evidence established by mineral caf e t e r i a s (Stockstad, 1953) showed that a l l sodium compounds received a much greater degree of use by ungulates than did any other compound. It also seems s i g n i f i c a n t that cobalt compounds were used to a degree second only to sodium. These two elements, as stated, have not been shown to be important i n plant growth. Chloride 80 compounds, other than sodium compounds, received a minor amount of use. The re s u l t s of this study are shown i n Tables 10 and 11. S i m i l a r l y , preference tests c a r r i e d out by B i s s e l (1953)? showed sodium to be the important element. Of 8 stakes con-taining sodium compounds 6 were used excessively, giving a 75 percent index of usage. Stakes containing other compounds were not used at a l l . Table 10 Comparative use by big game animals of mixtures offered i n sixteen mineral cafeterias i n western Montana, for a two year period (1951-52). From Stockstad (1953). No. of No. of T o t a l Compounds Used Cafeterias Cafeterias Amount of Amount of In The i n which In Which Frequency Mixture Mixture Percentage C a f e t e r i a The Mixture Mixture Of Use As Offered Used Of Mixture Relative Mixtures Was Offered Was Used Percentage In Pounds In Pounds Consumed Use Index NaHC03 9 9 100.0 21 19.1 91.0 l c 9 l Nal 16 16 100.0 .56 ^6.0 82A 1.82 NaCl 16 16 100.0 56 hOB0 71.5 1.72 NaH 2P0k 16 16 100 oO 56 "+L1 62;-3 1.62 CoCl2 36H 20 16 5 31.2 56 2,6 5.0 .36 KC1 16 h 25.0 56 2,0 3.6 .29 MgCl2°6H20 16 3 . 18.7 56 1,7 3.0 .22 (NHi^HPOif 6 1 16.7 26 .7 3 .0 .20 CaCl2 ° 6H 20 16 2 12.5 56 1.5 2 .7 .15 H3P0l+ KH 2P0l f 5 Ci 0 8 0 0 0 5 0 0 1*+ 0 0 0 Mgo(P0lf)2oI+-H20 1 0 0 2 0 0 0 NHLC1 16 0 0 12 0 0 0 FC1 >+ 0 0 9 0 0 0 C U S O L L 16 0 0 56 0 0 0 FeS0j«>7Hp0 lk 6 0 56 0 0 0 HoSOk (NH^SO^ 15 0 0 16 0 0 0 8 0 0 28 0 0 0 Calp 5 0 0 11 0 0 0 Mgl 2 0 0 6 . 0 0 0 5 0 0 11 0 0 0 KHCOO h 0 0 6 0 0 0 Control 16 0 0 56 0 0 0 co Table 11 Comparative use by big game animals of f i v e s o i l impregnation tests i n western Montana, for a two year period (1951-52). From Stockstad (1953). Compounds Used For S o i l Impregnations No, of Times Number of Frequency Compound Times Compound Of Use as a Was Offered Was Used Percentage T o t a l Pounds of Treated S o i l Consumed (Approx.) NaCl Nal NaH2P01f K C 1 M g C l 2 - 6 H 2 0 C O C I 2 . 6 H 2 Q C a C l 2 . 6 H 2 0 CuSO).. FeS0lf .7H 2 O HpSOk (NHl+)2HP01f H O P O L L Mg3(P0J+) 2. 1+-H20 (NHl f) 2SCi f Percentage of Volume Used Relative Use Index 16 15 93.7 ^36 100.0 1.9^ 18 16 89.0 *+03 92.5 1.82 13 7 53.8 176 hO.h 18 10 55.5 50 11.5 .67 10 2 20.0 6 ,21 18 X 5.6 3 .7 .06 18 0 0 0 0 0 18 0 0 0 0 0 16 0 0 0 0 0 9 0 0 0 0 0 2 0 0 0 0 0 3 0 0 0 0 0 1 0 0 0 0 0 6 0 0 0 0 0 CO ro 83 Packard (19^ -6) studying bighorn sheep, put two types of s a l t blocks onto the range: one contained a combination of calcium, phosphorus, sodium chloride and iodine s a l t s while the other was a block of sodium chloride. The sodium chloride block was consumed more r a p i d l y . Cowan and Brink (l91+9) conclude that magnesium and calcium are present i n such abundance i n the native rock of the region i t i s inconceivable that the vegetation i s lacking i n these elements. S i m i l a r l y , iron i s widespread i n these s o i l s . However, evidence suggests that boron i s not e s s e n t i a l i n ungulate n u t r i t i o n . Work by Gordon et. a l . (195*+) has shown that phos-phorus d e f i c i e n t c a t t l e w i l l select phosphorus when offered a choice of minerals. Since phosphorus was not selected i n the preference tests described previously, i t i s not l i k e l y that phosphorus i s a t t r a c t i n g goats to the l i c k s . Demarchi (1965) working i n the Ashnola region of B r i t i s h Columbia gives the calcium-phosphorus r a t i o for 5 species of grass. Samples col l e c t e d i n March contained the lowest concentrations of both calcium and phosphorus while those c o l l e c t e d i n the f a l l con-tained the highest concentration and more nearly approximated the r a t i o of 2:1, Dietz et. a l . (1962) working i n Colorado, showed the phosphorus concentration to be highest i n late spring with the calcium-phosphorus r a t i o c l o s e l y approximating 2 s i at t h i s time. In the f a l l the calcium concentration i n -creased, varying the CasP r a t i o 0 McLean et, a l . (1963) working i n the i n t e r i o r of B r i t i s h Columbia showed that phosphorus was highest i n the early spring and adequate for maintenance. The phosphorus l e v e l of feed, for the rapid growth of yearling c a t t l e , i s .2 percent as suggested by the National Research Council. Concentrations i n the Ashnola bighorn range i n March are much below this5 those i n Colorado i n the early spring are much higher and those c o l l e c t e d by McLean et. a l . (1963) i n July are s l i g h t l y lower. McLean and Tisdale (i960) showed that phosphorus was much above the rapid growth re-quirements for yearling c a t t l e , i f concentrations were de-termined i n May and June. This evidence would suggest that phosphorus i s extremely low at the end of winter but that the new growth i n early spring ( e s p e c i a l l y the leaves) i s extremely high i n phosphorus. It has been shown that goats usually feed on the spring vegetation at low elevations before using the l i c k s . It does not seem l i k e l y that phosphorus i s a t t r a c t i n g them to the l i c k s after they have fed on vegetation containing the highest yearly phosphorus concentrations. According to G i l b e r t (1957) potassium i s one of the plant macronutrients that i s somewhat less valuable to animals. Since i t forms a high proportion of the ash of plants as shown in Tables 15 and 16, i t should not cause any d e f i c i e n c i e s i n 85 the animal. The C a l i f o r n i a F i s h and Game Department ( B i s s e l , 1953)» concluded i n their preference tests that chlorine was not the preferred element since sodium compounds with chloride and bicarbonate were selected with equal frequency. Cowan and Brink (19^9) found extremely low concentrations of chlorine i n the l i c k and conclude that t h i s small amount probably i s not at t r a c t i n g the animals to the l i c k s . In the preference tests c a r r i e d out by Stockstad (1953) copper and s u l f a t e compounds received no use. It has not been shown that animals d e f i c i e n t i n manganese, zinc, molybdenum or copper w i l l select forages containing high per-centages of these elements. Since the d e f i c i e n c i e s i n the animal do not appear to be extreme, (no gross symptoms) i t i s not l i k e l y that these elements, i f s l i g h t l y d e f i c i e n t , would induce a craving i n the animal. Through successive elimination, most elements, with the exception of sodium and cobalt have been shown to be unimportant i n a t t r a c t i n g animals to the l i c k s . It has been shown by Wallace, Toth and Bear (191+7) i n their examination of some New Jersy plants that sodium can be d e f i c i e n t i n the vegetation. With the exception of a few accumulator plants, most of the 300 plants they examined were extremely low i n sodium. Most species of grass ranged from ,00 to .09 percent 86 for sodium. Other authors, notably* Harmer and Benne (19^-5), Collander (19*+1) and Lehr (19*4-1) have shown that sodium appar-ently has no s p e c i a l function i n plants. Plants benefit from i t only i n a s i t u a t i o n of potassium deficiency. Huffaker and Wallace (1959) state that differences i n Uptake capacity between plant species seemed more important than potassium levels i n determining the amount of sodium found i n the a e r i a l parts of plants. Potassium l e v e l appeared more important than plant species differences i n determining the amount of sodium i n the roots. There are a few exceptions to the above statements, as shown by sugar beets and some plants from saline areas, which may accumulate sodium. Determination of major forage species of the goat During the summer goats may be found over a wide range of elevations which includes a l l major vegetation zones i n t h i s area. The major forage species consumed at this time were determined using rumen samples and two confirming studies i n adjacent areas. The analysis of rumen samples, shown i n Table 12, suggests that grasses and sedges form a major part of the goat's diet at this time. Other studies suggest that the goat i s a snipfeeder and w i l l consume a large v a r i e t y of forage types. The study of Cowan (19Mf) working i n Banff and Jasper National Parks north of my area, and that of Saunders 87 Table 12 Rumen analyses from f i v e mountain goats, showing the percentage of forage types. ALL SAMPLES SHOT IN THE FALL OF I965 I. FEMALE - LAZY LAKE LICK - ADULT 1, Grass - 6.1.5$ 2. Browse - 3 8 .W Trace: Rock, F i r Needles I I . FEMALE - WILDHORSE CREEK - ADULT 1. Grass - 98$ 2. Browse - 2% I I I . MALE - ELK CREEK LICK - YEARLING 1. Grass - 53.3$ 2. Browse - k6.6% Trace: F i r Needles, and grouseberry IV. BULL RIVER AREA - ADULT MALE 1. Grass - 69% 2. Shrubs - 25% 3. Forbs - 6% V. FIND LAY CREEK AREA - ADULT FEMALE 1. Grass -2. Shrubs - 12% No conifer needles A l l animals shot i n alpine or subalpine areas 88 (1955)? working south of the study area i n Montana, l i s t the major or preferred forage species for d i f f e r e n t seasons of the year. Using these two studies, forage species were coll e c t e d i f l i s t e d as preferred by Cowan or Saunders and i f present i n the study area. Some species were present i n regions 1, 2 and 3 while others were r e s t r i c t e d to only one region. The species composition varied from one s i t e to the next showing the effects of elevation. In essence, the forage species c o l l e c t e d represent a composite picture of the major forage species present i n the area. Since neither s e l e c t i v i t y t r i a l s nor a complete food habits study could be ca r r i e d out, i t was hoped that analysis of the major forage species l i s t e d by other authors would be adequate and serve as an i n d i c a t i o n of the sodium content of the vegetation. Mineral content of the forage The forage species c o l l e c t e d were analyzed for t o t a l ash, sodium'-'and potassium. The r e s u l t s are shown i n Tables 1 3 , l ^ , 15 and 1 6 . The majority of forage species had sodium con-centrations i n the range of , 0 0 1 percent to . 0 0 7 percent, while the majority of potassium concentrations were between . 5 and 1.5 percent. This gives an average sodium-potassium r a t i o of l s 2 5 0 . The sodium values were i n general, extremely low^ much lower than any reports found i n the l i t e r a t u r e . The potassium values appeared adequate for normal health and growth of the 89 plant since no gross symptoms of potassium deficiency i n the plants were noted, G i l b e r t (1957) reports that approximately .2 to .6 percent potassium i n the dry matter of feed i s re-quired for animals. The concentrations i n the forage of the study area appeared to meet these requirements. Four samples were found to have s l i g h t l y higher sodium values and are as followss alpine bluegrass c o l l e c t e d i n September from region 1, .0V5 percent; alpine strawberry c o l l e c t e d i n September from region 1, .013 percent5 north-western sedge c o l l e c t e d i n the spring from region 1, .01*+ per-cent} Saskatoon berry c o l l e c t e d i n September from region 3? .016 percent. F i e l d sampling procedure could account for t h i s v a r i a t i o n since the same species c o l l e c t e d at other elevations and times do not show these higher concentrations. Other species c o l l e c t e d i n the same l o c a l area as these do not have such high concentrations. This may suggest that these four species may accumulate sodium where i t i s a v a i l a b l e . In gen-e r a l , the potassium concentrations tended to be lower i n proportion to the sodium concentration of these four species. The sodium-potassium r a t i o of the four species i s Is 11. The average potassium concentration i s .52 percent. This coincides with information obtained by Harmer and Benne (19*+5) who suggest that i n plants, the absence of sodium apparently r e s u l t s i n an increased absorption of potassium. T a b l e 13 F o r a g e s a m p l e s c o l l e c t e d i n S e p t e m b e r , 1966, i n r e g i o n s 1, 2 and 3 a t t he Toby-C r e e k l i c k , s h o w i n g t h e amount o f a s h as a p e r c e n t a g e d r y w e i g h t . A s h as Ash a s A s h as a % D r y a % D r y a % D r y R e g i o n 1 W e i g h t R e g i o n 2 W e i g h t R e g i o n 3 W e i g h t S t r a w b e r r y 7.7 S t r a w b e r r y 5.2 A l p i n e F i r 2.^ D o u g l a s F i r 1.2 D o u g l a s F i r 1.6 G r o u s e B e r r y 2.3 G r o u s e B e r r y W i l l o w h.5 W i l l o w 5.1 A l p i n e B l u e g r a s s 13.3 N o r t h w e s t e r n Sedge 5.5 N o r t h w e s t e r n Sedge I d a h o F e s c u e * K 3 I d a h o F e s c u e 11.3 T h i c k s p i k e W h e a t g r a s s B l u e b u n c h W h e a t g r a s s B l u e b u n c h W h e a t g r a s s P i n e G r a s s 9.1 P i n e G r a s s 13.1 P i n e G r a s s 1^.9 S a s k a t o o n B e r r y 5.2 S a s k a t o o n B e r r y 5.8 S a s k a t o o n B e r r y 13.3 B e a r B e r r y .7 Bea r B e r r y 3.0 B e a r B e r r y 2.1 As pen 5.1 A s p e n h.H A s p e n ^.2 B u f f a l o B e r r y B u f f a l o B e r r y 6.0 S p i k e T r i s e t u m h.h C a r e x S p p . 3.h S u b a l p i n e B e e d l e g r a s s R i c h a r d s o n N e e d l e g r a s s R o c k y M t . J u n i p e r 5.2 R o c k y M t • J u n i p e r K e n t u c k y B l u e g r a s s 7.1 vO O Table 1*+ Forage samples c o l l e c t e d i n the spring, 1966, i n regions 1, 2 and 3 at the Toby-Creek l i c k , showing the amount of ash as a percentage dry weight. Ash as Ash as~ Ash as a % Dry a % Dry a % Dry Region 1 Weight Region 2 Weight Region 3 Weight Strawberry 5.2 Alpine F i r 2.2 Douglas F i r 2.2 Douglas"" F i r " 2.2 Grouse Berry 3.2 Grouse Berry 6.H-Willow 3.1 Alpine Bluegrass 5.3 6.2 Northwestern Sedge 8.0 Northwestern Sedge Northwestern Sedge 6.8 Idaho Fescue 6.6 Idaho Fescue 11.1 11.6 Bluebunch Wheatgrass 11.0 Pine Grass ?•? Pine Grass Pine Grass 10.3 Saskatoon Berry K\ Saskatoon Berry 6.2 Saskatoon Berry if.6 Bear Berry 2.^ Bear Berry 2.6 Bear Berry 1.8 Aspen h.h Aspen 7.0 Aspen 7.5 Buffalo Berry Kl Buffalo Berry ^.6 Richardson Needlegrass 9.9 June Grass 8.9 Carex spp. 7.0 vO H Table 15 Forage samples c o l l e c t e d i n September, 1966 i n regions• 1, 2 and 3 at the Toby Creek l i c k , showing the sodium and potassium concentrations as a percentage dry weight. Region 1 Strawberry Alpine F i r Grouse Berry Willow Alpine Bluegrass Northwestern Sedge Idaho Fescue Thickspike Wheat-grass Pine Grass Saskatoon Berry Bear Berry As pen Buffalo Berry Spike Trisetum Carex spp„ % Na % K Region 2 % Na % K Region 3 .0129 068 .0026 .61 Douglas F i r .0030 o0026 M .0055 .75 .0^5 .67 .001^ o38 .0023 .52 Bluebunch Wheat-O0036 .86 grass .0015 0OOI3 .5^ Pine Grass .0026 o00^9 1.17 Saskatoon Berry .0091 o0CA3 .53 Bear Berry .0035 .0073 .67 Aspen .0051 .0038 .70 o0021 .6*f oOO^fl .90 Subalpine Needle-grass .0056 Rocky Mt. Juniper .0071 Strawberry .57 Douglas F i r Grouse Berry Willow Northwestern Sedge Idaho Fescue Bluebunch Wheat-.36 grass .65 Pine Grass .86 Saskatoon Berry ..V3 Bear Berry .90 Aspen Buffalo Berry Richardson Needle-,71 grass ,k8 Rocky Mt. Juniper Kentucky Bluegrass June Grass % Na % K .0077 .81 .0017 .5^ .0011* .2^ .0036 .6*+ .0017 .29 .OOl^ .2*+ .0012 .30 .0021 .67 .016 .^6 .0031 M .0036 .65 .0055 lo00 .0030 .53 ,0033 .63 .0015 .36 .0038 .35 ro Table 16 Forage Samples c o l l e c t e d i n the spring, 1966, i n regions 1, 2 and 3 at the Toby Creek l i c k , showing the sodium and potassium concentrations as a percentage dry weight. Region 1 Strawberry Alpine F i r Grouse Berry V i l l o w Alpine Bluegrass Northwestern Sedge Idaho Fescue Pine Grass Saskatoon Berry Bear Berry Aspen Buffalo Berry Carex spp. % Na % K Region 2 % Na % K Region 3 .0071 1.07 .002M- .44 Douglas F i r .0028 ..»f 5 .0035 .83 .004-1 1.09 ,0lV .29 Northwestern Sedge .0015 -6M-.0032 1.00 Pine Grass .0066 .87 Saskatoon Berry .00V3 .36 Bear Berry .0045 1.09 Aspen .0049 1.23 ,0055 2.04 .0027 .0053 ,57 Douglas F i r Grouse Berry .if9 Northwester Sedge Idaho Fescue Bluebuhch Wheatgrass ,0060 2.36 Pine Grass ,0063 .91 Saskatoon Berry ,0085 .32 Bear Berry ,0051 1.41 Aspen Buffalo Berry Richardson Needlegrass June Grass % Na % K o0020 .55 .0035 .16 .0018 .35 .0028 8 ,0060 .77 .0032 1.14 .0039 .97 ,0023 .31 .0050 1.36 .OQifl 1,18 .OOifO .if 8 ,0020 .60 vO 94 In his food analysis study, Saunders (1955) gives alpine bluegrass the highest observed summer use frequency by the mountain goat. The high degree of use coincides with the high sodium concentration and s e l e c t i v i t y t r i a l s may prove i n t e r e s t i n g . Denton and Sabine (1961) have shown that an intake of 100 m. - equivalents of sodium d a i l y , led to a sodium craving i n sheep, while an intake of 500 m. - equivalents per day reduced the craving. If a goat were fed s o l e l y on alpine bluegrass with a concentration of .045 percent x i t would re-quire 51.1 grams of the grass to supply 100 m. - equivalents of sodium per day and 255.5 grams to supply 500 m. - equiv-alents per day. This represents an almost impossible intake of alpine bluegrass i n one day. Neither the d i s t r i b u t i o n of t h i s grass, nor i t s abundance i n the study area i s known. Since a l l other species are at least 7 times lower i n concen-t r a t i o n , i t i s not l i k e l y that the sodium requirements of the goat are being supplied by the vegetation. It has been shown that a l l plants c o l l e c t e d i n the spring and f a l l , from regions 1, 2 or 3, have extremely low sodium values. Since the l i c k has been described as a runoff or accumulation s i t e , i t was thought that plants growing at higher elevations would have a lower sodium and ash content. As shown i n Figures 18 and 19 the sodium concentration does X o *i J0030 Q ui O < .0020 z U l V 2 0010 0. < o — .0070 < 2 .0060 ^ . 0 0 5 0 O U .0040 2 003G Q O «n 0020 0010 Fir Saskatoon Berry 2 Region .0060 .0050 .0040 0O3O .0020 J0010 0090 .0080 .0070 0060 .0050 .0040 .0030 0020 OOIO Northwestern Sedqe Bear Berry 2 Region .0060 .0050 .0040 .0030 .0020 .0010 Pine Grass .0060 XJ050 .0040 .0O30 OO20 .0010 Aspen 2 3 Region FIGURE 18. A comparison of forage samples collected in the spring showing sodium concentration plotted against elevational region. .ooiq 1 2 3 1 2 3 1 2 3 Region Region Region FIGURE 19. A comparison of forage saMples collected in September showing sodium concentration plotted against elevational region. X o m  1 o < u et Ul a. < i/) < I < Douglas Fir Buffalo Berry Saskatoon Berry Northwestern Sedge 12 10 8 6 4 Bear berry Pine Grass 12 10 8 6 41 Grouseberry Idaho Fescue 1 2 3 1 2 3 1 2 3 1 REGION FIGURE 20. A comparison of forage samples collected in the spring showing ash as a percentage dry weight plotted against elevational region. REGION 21. A comparison of forage samples collected in September showing, ash as a percentage dry weight plotted against elevational region. 95 not appear to be consistently higher i n region 3 i n either spring or f a l l . In Figure 21, i t can be seen that the ash content i s not consistently higher i n region 3, during the f a l l . In the spring, no consistent trend could be found as shown i n Figure 20. Plants c o l l e c t e d i n the spring and f a l l d i f f e r e d i n the state of maturity. The spring sample consisted of plants i n an early growth stage, containing a high proportion of water. B l a i r and Epps (1967)> report that rusty blackhaw (Viburnum rufidulum) was highly succulent i n the spring, with leaves averaging 79 percent moisture and twigs 76 percent. As shown i n Figures 22 and 23, the shrubs and trees showed no consistent trend i n sodium values from the spring to f a l l . The grasses and sedges c o l l e c t e d i n region 1, as shown i n Figure 25 do not show any consistent trend from spring to f a l l . As shown i n Figure 24, grasses and sedges c o l l e c t e d at the l i c k are consistently higher i n sodium i n the spring than i n the f a l l . The great degree of v a r i a t i o n evidenced here can well be attributed to s o i l , s i t e f a c t o r s , s o i l moisture and vege-t a t i o n type. According to Cook and Harris (1950) the v a r i a t i o n i n chemical composition of plants i s affected by the v a r i a -b i l i t y of botanical composition of the range, stage of growth, available s o i l moisture, temperature, s o i l types, s i t e and X o a m .0030| o P- 0020 Z Ui U .0010 et .020 0. < < z o < et Z joio ui U .0090 O 0080 u <g .OO70 2 0 0 6 0 a O 0 0 5 0 in .0040 Douglas Fir Fall Spring Saskatoon Berry Fall Spring .0060 0050 .0040 0030 0020 0010 Aspen 0O4O 0O3O .0020 0010 Grouseberry Fall Spring Buffalo Berry Fall Spring FIGURE 22. A comparison of spring and fall samples from region 3 using sodium concentration as a percentage dry weight. z o a © < u ce, in a. . 0 0 3 0 . 0 0 2 0 . 0 0 1 0 < < z . 0 0 7 0 o ATI . 0 0 6 0 i-Z . 0 0 5 0 U l VJ z . 0 0 4 0 o u 2 . 0 0 3 0 3 O o . 0 0 2 0 l/ l . 0 0 1 0 Fall FIGURE 23. Alpine Fir Spring . 0 0 6 0 . 0 0 5 0 . 0 0 4 0 . 0 0 3 0 . 0 0 2 0 . 0 0 1 0 . 0 0 6 C M . 0 0 4 C M . 0 0 2 0 Fall TIME Willow Fall Spring Bearberry Spring J 0 0 3 0 . 0 0 2 0 . 0 0 1 0 . 0 0 5 0 . 0 0 4 0 O O 3 0 . 0 0 2 0 . 0 0 1 0 Grouseberry Fall Spring Buffalo Berry Fall Spring A comparison of spring and fall samples from region 1 using sodium concentration as a percentage dry weight. SODIUM CONCENTRATION AS A PERCENTAGE DRY WEIGHT Sedg Fall Spring Fall Spring Fall TIME FIGURE 25. A comparison of spring and fall samples from region 1 using sodium concentration as a percentage dry weight. 96 general climatic conditions 0 It i s not within the scope of t h i s study to attribute the v a r i a t i o n i n ash and sodium con-tent to s p e c i f i c , environmental or b i o l o g i c a l factors but only to show where trends may or may not e x i s t . 97 LICK SOILS Introduction In general, the use of natural earth l i c k s by ungu-lates i n the Rocky Mountain Trench of B.C. i s confined to the summer months. At th i s time goats appear to prefer dry earth l i c k s but on occasion v i s i t mineral springs. Several other big game species u t i l i z e the l i c k s i n the study area during the early spring and summer but use by goats i s predominant. A l l l i c k s are composed of a dry whitish clay but each contains a d i f f e r e n t amount of sand and gravel. It has been shown that i f two l i c k s are adjacent to one another i n a c e r t a i n area, goats usually prefer one over the other. In most cases, the preferred l i c k has less sand and gravel. With-i n an earth l i c k , s i t e s are chosen by the goat to which i t returns frequently to eat the s o i l . These high l i c k i n g s i t e s make up about 1 to 5 percent of the t o t a l surface area of the l i c k but approximately 90 to 95 percent of the actual l i c k i n g occurs here. In most cases, these pockets of highly preferred s o i l occur among the roots of Douglas f i r trees or i n i s o l a t e d spots i n the l i c k . High l i c k i n g s i t e s at the Lazy Lake and Mary Creek l i c k s are shown i n Figures 26 and 27. Where the high l i c k i n g s i t e s occurred on a bank, i t -was seen that they were i n the "B" horizon or accumulation region of the s o i l p r o f i l e . In some cases the l i c k i n g s i t e s FIGURE 27. A high licking site at the Mary Creek lick. 9 8 extended into the lower region of the "A" layer«, In these preferred s i t e s the clay was usually dark brown and moist,, Williams (1962) suggests that the character of the s o i l seemed to be a p r i n c i p a l determinant of the quantity of s o i l consumed at a particular l i c k . He noticed that l i c k s within the same s a l t ground would not always be used to the same degree. In the regions of heaviest elk use he states that a strong pref-erence was shown for the more recently established s a l t l i c k s . Consequently, c a t t l e s a l t grounds that were constantly being replenished with s a l t were often used the heaviest. Salt l i c k analyses c a r r i e d out by the C a l i f o r n i a F i s h and Game Department ( B i s s e l , 1953) showed that i n every i n -stance sodium was higher i n the l i c k than i n the con t r o l , Beeman (1957) found that the sodium concentration of the av-erage l i c k was 16 times greater than the contents found i n the water from 3 other l i c k s , l i t t l e used by animals. High and low l i c k i n g s i t e s As has been shown, c e r t a i n sites i n a l i c k receive more use than do others. Closer examination revealed the following physical differences? high l i c k i n g s i t e s contained a large proportion of moist brown clay and were usually among the roots of Douglas f i r trees, the greatest amount of l i c k i n g was usually i n the "B" horizon of the s o i l p r o f i l e 5 the low l i c k i n g s i t e s were composed of dry, grey clay, contained more FIGURE 26. A high licking site at the Lazy Lake lick, among the roots of a Douglas Fir tree. A rattrap marker is shown in the foreground. FIGURE 27. A high licking site at the Mary Creek lick. 99 sand and rocks and were not situated beneath f i r trees. At the Lazy Lake l i c k about 95 percent of the high and low l i c k i n g s i t e s are beneath Douglas f i r trees. Those s i t e s which are the deepest, contain more shale rock and appear the oldest, are the least preferred. The preferred s i t e s contain less shale, more available s o i l and are not as deep. At the Dutch Creek l i c k observations showed that the animals moved i n a pattern while using the l i c k . There are 3 groups of high l i c k i n g s i t e s which form a t r i a n g l e . Each group i s about 100 feet apart. On August 15, a group of 9 animals entered the l i c k . There were three females and three kids, a female without a kid and two yea r l i n g s . The animals moved from one high l i c k i n g s i t e to another, with occasional use of low l i c k i n g s i t e s when the preferred s i t e s were being used by other animals or as they moved from one preferred s i t e to another. The females with young dominated the high l i c k i n g s i t e s and could displace a female without young or a yearling from i t by charging them. The yearlings occupied the high l i c k i n g s i t e s least often. It appeared that larger females could displace smaller ones from the high l i c k i n g s i t e s i f both had young. Mineral content The mineral content of 3 l i c k s was determined as shown i n Table 17. At the Dutch Creek l i c k , samples from high 100 and low l i c k i n g s i t e s were obtained from a primary and second-ary l i c k . The sodium concentrations of two high l i c k i n g s i t e s from the primary l i c k (samples 5 and 7) are consistently higher than the sodium concentrations from the low l i c k i n g s i t e s i n the same l i c k (sample 11). Samples from high l i c k i n g s i t e s i n the secondary l i c k (samples 8 and 1*0 are again higher than those from low l i c k i n g s i t e s (sample 12) i n the l i c k . Samples from high l i c k i n g s i t e s of the primary l i c k are higher than those from similar s i t e s i n the secondary l i c k . At. Toby Creek, a similar trend i s apparent, upon comparison of samples 10 and 15 to sample 9» Sample 10 was co l l e c t e d at the base of the MB" layer i n the s o i l p r o f i l e and sample 15 near the top of this layer." Sample 10 could almost be considered a low l i c k i n g s i t e due to i t s position i n the p r o f i l e . The sodium concen-t r a t i o n c l o s e l y approximates that of sample 9» At Lazy Lake a s p e c i a l sample was taken from a high l i c k i n g s i t e . This consisted of a white encrustation on the surface of the s o i l where goats spent much time l i c k i n g . As shown i n Table 17 the sodium concentration of thi s sample (number 16) was at least 10 times higher than that i n other l i c k s . The calcium and phosphorus content was also much higher. The cobalt was lower than the majority of samples from other l i c k s . On examining calcium and cobalt concentrations from high and low l i c k i n g , i t does not appear that a similar trend 101 e x i s t s as t h a t f o u n d f o r s o d i u m . P h o s p h o r u s , as d e t e r m i n e d f o r t h a t a v a i l a b l e t o p l a n t s was n o t f o u n d i n amounts s u f f i c i e n t t o a t t r a c t a n i m a l s . New l i c k s D u r i n g t h e two summers o f f i e l d work i t became o b v i o u s t h a t new l i c k s i t e s were b e i n g c r e a t e d . At L a z y Lake two new l i c k s i t e s were f o u n d d u r i n g t h e summer o f 1966. I t was known t h a t one l i c k h a d n o t b e e n t h e r e i n t h e f a l l o f 196U-. B o t h s i t e s were c l o s e t o the m a i n l i c k . One was a p p r o x i m a t e l y two f e e t s q u a r e and one f o o t d e e p , t h e o t h e r s l i g h t l y s m a l l e r . S i n c e t h e y were b o t h i n t h e v i c i n -i t y o f t he m a i n l i c k i t seemed p r o b a b l e t h a t new l i c k s s h o u l d be s t a r t e d b e c a u s e o l d l i c k s become c l o g g e d w i t h s h a l e r o c k and r o o t s o f D o u g l a s f i r . S i n c e d e e r , s h e e p and g o a t s were a l l p r e s e n t i n t h i s v i c i n i t y i t i s n o t p o s s i b l e t o d i s t i n g u i s h t h e r o l e e a c h s p e c i e s p l a y e d i n t h e i r f o r m a t i o n . A t D u t c h C r e e k a u n i q u e s i t u a t i o n o c c u r r e d . D u r i n g the summer o f 1965 a l o g g i n g o p e r a t i o n was u n d e r t a k e n a b o u t two m i l e s b e l o w t h e l i c k on t h e g u i d e t r a i l w h i c h r a n f r o m t h i s s i t e p a s t t h e l i c k . T h i s o p e r a t i o n was d i s c o n t i n u e d a t t h e end o f t h e summer i n 1965? a f t e r t h e g o a t s had c e a s e d u s i n g t h e m a i n l i c k . Many s k i d t r a i l s h a d b e e n e x t e n d e d t o w i t h i n one m i l e o f t h e l i c k . A r o u n d t h e end o f June d u r i n g t h e summer o f I966, g o a t h a i r was f o u n d on t h e g u i d e t r a i l b e l o w t h e m a i n Table 17 Chemical analyses of 3 l i c k s , comparing high and low l i c k i n g s i t e s 0 NHUAc. pH Sample No. Description Na , PH ° 7, (p.p.m.) Ca • 0 Co Na 2.0 207 N.D. 2.0 185 N.D. ikQ 2.0 160 2.0 160 N.D. 115 2.0 115 N.D. 160 N.D. N.D. ^,950 Available P (p.p.m.) D 5 Good s i t e (1) D 6 Skid t r a i l D 7 Good s i t e (1) D 8 Good s i t e (2) T 9 Poor s i t e T 10 Good s i t e D 11 Poor s i t e (1) D 12 Poor s i t e (2) D lk Good s i t e (2) T 15- Good s i t e L 16 Good s i t e 185 1+38 207 169 160 160 138 115 160 ' 3^ -5 5,500 10,000 10,^00 11,000 1^ 050 8,^00 8,000 11,000 10,^00 12,000 7,200 H-7,000 21^00 85,000 2^,000 1,050 29,^00 29,200 21,H-00 18,600 25,000 22,600 62,000 3.0 5.0 2 h.O h:o h.o 2.0 ^.0 3.0 2.0 0 Primary 0 0 Primary 0 Secondary 0 PrImary 0 Primary 0 Primary 0 Secondary 0 Secondary 0 Primary 25 Primary N.D. - Not Determinable D - Dutch Creek l i c k T - Toby Creek l i c k L - Lazy Lake l i c k (1) - Main l i c k (2) - Secondary l i c k H O ro 103 l i c k and o n t h e s k i d t r a i l s , Many f r e s h t r a c k s and much f e c a l m a t e r i a l was a l s o p r e s e n t , , I t a p p e a r e d t h a t some g o a t s , u p o n r e a c h i n g t h e m a i n l i c k c o n t i n u e d down t h e g u i d e t r a i l t o t h e s k i d t r a i l s where l i c k i n g was d o n e . E v i d e n c e o f l i c k i n g was p r e s e n t on t h e t h r e e t o f o u r f o o t banks o f t h e s k i d t r a i l s . S i n c e g o a t s were n o t b e l i e v e d t o have e v e r moved down t h i s l ow b e f o r e i t l e d t o many s p e c u l a t i o n s . M i n e r a l c o n t e n t o f t h e m a i n l i c k was c o m p a r e d t o m i n e r a l c o n t e n t o f t h e e a r t h f r o m t h e s k i d t r a i l s as shown i n T a b l e 17. G o a t s a r e v e r y s p e c i f i c i n t h e t r a i l s t h e y u s e and d e v i a t i o n f r o m them i s l i m i t e d . I t i s c o n c l u d e d t h a t g o a t s wander t h r o u g h t h e t i m b e r v e r y l i t t l e . I t i s n o t t h o u g h t t h a t t h e y b e g a n u s i n g t h e s k i d t r a i l s by a c c i d e n t b u t by u s i n g a d e f i n i t e t r a i l came t o a c q u i r e a new l i c k s i t e . The c o m p a r i s o n o f a n a l y s e s shows t h a t t h e s o d i u m c o n c e n t r a t i o n o f t h e s k i d t r a i l s i s much h i g h e r t h a n t h e h i g h l i c k i n g s i t e s i n t h e p r i m a r y l i c k . T h i s i s i n a g r e e m e n t w i t h the s t a t e m e n t o f Beeman (1957) made e a r l i e r i n t h i s s e c t i o n . At t h i s t i m e , g o a t s have h a d a c c e s s t o t h i s new l i c k f o r o n l y one summer . D u r i n g t h e summer o f 1967 i t w i l l b e i n t e r e s t i n g t o s ee i f more g o a t s u s e t h i s l i c k o A t D u t c h C r e e k , t h e s e c o n -d a r y l i c k i s more a c c e s s i b l e t o g o a t s e n t e r i n g f r o m r e g i o n 1, t h e p r i m a r y l i c k i s l e s s a c c e s s i b l e and t h e s k i d t r a i l s a r e i n t i m b e r , l i t t l e t r a v e l l e d by g o a t s . The s o d i u m c o n c e n t r a t i o n and d e g r e e o f u s e by g o a t s i s h i g h e s t i n t h e s e l a t t e r two l i c k s . 10k The selectiveness shown by goats for s i t e s high i n sodium leads to the conclusion that sodium i s the attra c t i n g element. 105 SODIUM IN THE ANIMAL Introduction The e f f i c i e n c y of the various homeostatie mechanisms i n the animal body i s very h i g h o The regulatory functions o f the r e n a l mechanisms, the role of the adrenal hormone; aldo-sterone , i n minute quantities, i s a most important regulator of sodium balance, the transfer of water between c e l l s and ex t r a c e l l u l a r f l u i d , the r o l e of bone i n mobilizing sodium i n response to acute acidosis and s a l t depletion and the role of the c e n t r a l nervous system and i t s influence on sodium met-abolism a l l contribute to t h i s e f f i c i e n c y (Forbes, 1962), He feels that serum sodium remains constant under conditions of a sodium load which i s excreted promptly or i n the case of a sodium deficiency, sodium excretion drops to a very low l e v e l within two to four days aft e r removal of sodium from the d i e t . Certain conditions are associated with a disordered sodium metabolism. When homeostatie mechanisms f a i l because of disease, or when the i r regulatory l i m i t s are exceeded by the magnitude of the stress put upon them,, sodium metabolism i s upset. Some conditions which may lead to disordered sodium metabolism ares extreme g a s t r o i n t e s t i n a l losses, urinary losses associated with r e n a l i n s u f f i c i e n c y , adrenocortical f a i l u r e s a c e n t r a l nervous system disease and extreme sweating. These conditions leading to sodium depletion usually r e s u l t i n 106 c l i n i c a l symptoms such ass a decrease i n serum sodium, a de-crease i n plasma volume, an increase i n the hematocrit value. Shock and eventual death may r e s u l t (Forbes, 1962). It has not been shown that any of the above disorders, usually associated with the Individual, are common to pop-ulations of ungulates using natural earth l i c k s . Since gross c l i n i c a l symptoms have not been reported for animals using l i c k s , i t i s l i k e l y that the homeostatic mechanisms are functioning normally i n preventing a severe s a l t depletion, Marriott (1950) says that i f only intake of food ceases and water continues to be taken, i t i s a considerable time, In the absence of abnormal s a l t losses such as those previously des-cribed, before serious s a l t depletion occurs. During the summer of 1966, goats entering the Toby Creek l i c k were observed to have diarrhoea. The material they were passing was e n t i r e l y organic and no clay was found i n the droppings. This would indicate that diarrhoea occurred before they entered the l i c k , probably caused by the s h i f t from mature winter forage to succulent spring forage. Since diarrhoea Is a g a s t r o i n t e s t i n a l disorder, abnormal sodium loss to some degree could occur. According to both Marriott (1950) and Weisberg (1953)? urinary loss of sodium occurs for the f i r s t two to four days on a diet high i n water and extremely low i n sodium. It has been found by S e l l e r s and Roepke (195D that 10? extreme diuresis increased several fold,, the excretion cf calcium, sodium, chloride, and phosphate. During d i u r e s i s , the urine contained more of these e l e c t r o l y t e s as they were "washed out by the water," than urine c o l l e c t e d before d i u r e s i s 0 It would appear, that to a limited e x t e n t a b n o r m a l s a l t losses can occur\ that excessive water intake can remove sodium during the f i r s t two to four days and diuresis can "wash" sodium out of the body. The goat, as described, i s affected by these three factors prior to entering the l i c k . Serum sodium values obtained under these conditions, for the goat, ranged from 141 to 153 nu =• equivalents per l i t r e as shown i n Table 18, The s t r e s s f u l conditions of trapping and handling were not expected to a l t e r the normal sodium range as the l i t e r a t u r e does not suggest these factors as a f f e c t i n g serum sodium values. The serum sodium values obtained were comparable to those found i n domestic sheep (146 - 161) and c a t t l e (122 - 166) (Dukes, 1955), It was thought that the serum sodium values are probably within a normal range for the mountain goat 0 The value for the kid i s higher as i s the case for most young animals. According to Vrzgula (1965) the highest sodium, potassium and calcium values were found i n the serum of calves immediately after b i r t h . Mean values declined during growth and i n cows con-tinued to decline slowly but s i g n i f i c a n t l y throughout l i f e . 108 Table 18 Some serum sodium values and the packed-call volume obtained for the mountain goat. Class SeTMjm Sodium In m.eq/L. Packed C e l l Volume i n % Age i n Years Date Movement 1. Yearling male 153 1.5 May 26 To l i c k 2. Adult male 142 38 10, June 1 To l i c k 3. Adult male 151 41 5.5 June 12 To l i c k 4. Adult female (no young) 153 4.5 June 19 To l i c k 5. Adult female (with young) 141 3.5 July 1 Out of l i c k 6. Adult female (no young) A. 151 46 3.5 July 12 Out of l i c k B. 152 50 July 15 Holding pen c. 152 42 July 28 Holding pen 7. Adult male 153 6.5 July 19 To l i c k 8, Female k i d 160 3 months August Out of l i c k 109 Comparison of sodium values of animals moving into and out of the l i c k . Examination of a small sample of goats of both sexes (Table 18) revealed that there were no s i g n i f i c a n t differences i n serum sodium values between goats coming to the l i c k for the f i r s t time and those that had been using i t 0 Serum sodium values lay between l * f l and 153 m0 s q . / l i t r e and are thus within a normal range for healthy ungulates (Dukes, 1955). It i s known that serum sodium levels are maintained by some species even after they have developed a craving for the mineral as a consequence of dietary deficiency. For ex-ample, Smith and Aines (1959)? working with dairy c a t t l e , found that unsupplemented cows developed s a l t hunger i n about two weeks even though serum sodium values remained normals There was no difference i n serum sodium between supplemented and unsupplemented groups after 13 months, or between the f i r s t c o l l e c t i o n period at one month and the l a s t c o l l e c t i o n period at 13 monthso Beilharz ( I 9 6 3 ) working with sheep states that the appetite for sodium and plasma concentration can vary almost independently. A single attempt to a l t e r serum sodium values by feeding vegetation known to be sodium d e f i c i e n t , f a i l e d (Table 18). The hematocrit values, as shown i n Table ,18, range 110 from 38 percent to 50 percent, Rosen and Bischoff (1952) report a packed-cell volume of 4 4 . 8 - 0 . 5 percent for several sub-species of Odocoileus. Browman and Sears (1955) give the packed-cell volume of Rocky Mountain mule deer i n Montana as 39.6 percent. K i t t s et. a l , (1956) suggest that such factors as high a l t i t u d e , muscular exercjse, heightened environmental temperatures and c e r t a i n pathological conditions are known to cause an Increase i n the packed-cell volume. They state that the certainty with which such f i e l d data may be interpreted must rest largely upon the a v a i l a b i l i t y of normal values ob-tained from animals reared under known environmental conditions. He also suggests that the packed-cell volume increases with increasing age. Deer (Odocoileus) on a high plane of n u t r i t i o n had a packed-cell volume of 40 percent i n the 20 - 100 day age group and one of 58$ i n the 395 - ^65 day age group. Data co l l e c t e d on the mountain goat would suggest that i t s packed-c e l l volume i s within a normal range. Blood serum studies conducted on other animals show trends of c e r t a i n minerals i n the serum over one year periods. Stufflebeam (1964) states that Hereford cows which always got s u f f i c i e n t mineral and protein supplement showed no seasonal trend i n serum calcium and showed a sharp decline during winter of serum phosphorus. According to Payne (1964) inorganic phos-phorous was lower during l a c t a t i o n than i n the dry period and f e l l w i t h advancing age5 c a l c i u m was.not a f f e c t e d by l a c t a t i o n but f e l l s l i g h t l y w i t h age. Results were obtained from plasma s t u d i e s on the cow, V r z g u l a ( I 9 6 2) studying Pinzgou c a t t l e found that the lowest values f o r sodium & potassium and calcium were i n A p r i l . The highest values were i n the summer and late f a l l . He a t t r i b u t e s t h i s to the greater amounts of minerals and vitamins i n the feed, Denton (I96D found that when Merino sheep were o f f e r e d water and s o l u t i o n s of NaCl, NaHCO^? or KC1, they showed a preference f o r NaHCO^. Although t h e i r d i e t s u p p l i e d about 100 m, - e q u i v a l e n t s of sodium d a i l y , most sheep drank the s o l u t i o n s and increased t h e i r intake to 500 m. - e q u i v a l e n t s d a i l y . A study conducted by D e v l i n and Roberts (1963) on wether lambs f o r a period of 30 days showed that the concentrations of sodium, potassium and c h l o r i d e i n serum was the same on a l l d i e t s . The d a i l y intake of three d i e t s con-s i s t e d of Mf« and 129 ni. = e q u i v a l e n t s of sodium. I t has been shown that serum sodium i s r e g u l a t e d to a high degree i n the absence of abnormal c o n d i t i o n s . I t does not appear that sodium l e v e l s i n the blood change over short periods due to the c o n t r o l of homeostatie mechanisms and i t i s l i k e l y that the serum sodium values obtained-for the goat are normal. No d i f f e r e n c e could be found between animals entering and l e a v i n g the l i c k or among animals using the l i c k over the summer. From the preceeding section on vegetation i t i s known that phosphorus i n plants i s highest i n early spring and calcium gradually increases over the summer to reach a maximum i n late f a l l . The blood phosphorus declined during the winter and was lowest during l a c t a t i o n 0 Sodium appears to follow this pattern. Calcium does not appear to change as markedly,, This suggests that while the three elements may he reduced i n the animal during the spring, sodium i s the only one not r e a d i l y available from the vegetation,. 113 SUSPECTED SELENIUM DEFICIENCY During the summer of 1966, trapping operations brought many animals under stresses to which they were not accustomed. Animals were restrained with ropes, causing them to struggle -vigorously for varying lengths of time. The r e -su l t i n g gross symptoms c l o s e l y approximate those described by Muth (1963)? for a selenium d e f i c i e n t myopathy termed white muscle disease. Trapped animals, upon release, were hesitant or unable to r i s e to t h e i r feet so that many had to ba l i f t e d into a standing position. When i n th i s p o s i t i o n , a l l animals lacked c o n t r o l of the i r hind limbs and pelvic region and. showed reduced maneuverability. Recovery occurred for most i n about 5 minutes as they moved out of the trap. These symptoms were at f i r s t a ttributed to p a r t i a l loss of c i r c u l a t i o n . Three animals, two females and a k i d , which did not recover, were kept i n a holding pen a f t e r trapping. A l l d i e d following symptoms of l o s s of muscle c o n t r o l . Muscle p a r a l y s i s began i n the hind legs and pelvic region, moved to the front limbs and e v e n t u a l l y enveloped the e n t i r e body. During the period before death, the goats a t e , drank and defecated nor-mally. One female had extreme d i f f i c u l t y breathing just p r i o r to death, probably due to loss of c o n t r o l over the thoracic muscles. One female showed edema i n the hind l e g s . According to Rosenfeld and Beath (1964) edema i s common i n animals with w h i t e m u s c l e d i s e a s e . T h e y s t a t e t h a t s p o n t a n e o u s r e c o v e r y i s a l s o common and t h i s may e x p l a i n t h e s h o r t p e r i o d o f p a r a l y s i s e x h i b i t e d by t r a p p e d a n i m a l s . The k i d d i e d t h r e e weeks a f t e r t r a p p i n g , one. f e m a l e a f t e r two weeks and t h e o t h e r i n two d a y s . A u t o p s i e s d i d n o t r e v e a l a b n o r m a l i t i e s I n t h e a n i m a l s . As a r e s u l t o f t h e s e o b s e r v a t i o n s , 12 p l a n t s a m p l e s and 2 l i c k s a m p l e s were a n a l y z e d f o r me by M r . R . B . C a r s o n , o f t h e A n a l y t i c a l C h e m i s t r y R e s e a r c h S e r v i c e , i n O t t a w a , t o d e -t e r m i n e s e l e n i u m c o n c e n t r a t i o n s . The r e s u l t s o f t h i s a n a l y t -i c a l work a r e shown i n T a b l e 19. A l l t h e p l a n t s a m p l e s , w i t h t h e e x c e p t i o n o f b u f f a l o b e r r y , a r e i n t h e r a n g e b e l o w .1 p . p . m . s e l e n i u m . H i d i r o g l o u (I965) f o u n d n u t r i t i o n a l m u s c u l a r d i s e a s e ( w h i t e m u s c l e d i s e a s e ) i n b e e f c a t t l e where t h e v e g e -t a t i o n c o n t a i n e d l e s s t h a n .1 p . p . r n . A c c o r d i n g t o Ccwan and G u i g u e t (1965) t h e m o u n t a i n g o a t i s one o f t h e f e w a n i m a l s w h i c h w i l l e a t b u f f a l o b e r r y . The s amp l e c o l l e c t e d at t h e T o b y C r e e k l i c k c o n t a i n e d . 2 4 7 p . p . m . D u r i n g t h e summer o f 1966, on many o c c a s s i o n s , g o a t s were o b s e r v e d f e e d i n g on t h i s s p e c i e s i n t h e . v i c i n i t y o f t h e l i c k . It d o e s n o t a p p e a r that t h e c o n d i t i o n was a l l e v i a t e d due to s e l e c t i v e f e e d i n g on b u f f a l o b e r r y . Two s o i l s a m p l e s , one f r o m t h e T o b y C r a e k l i c k and one f r o m t h e D u t c h C r e e k l i c k a l s o c o n t a i n e d v e r y l i t t l e s e l e n i u m . It a p p e a r s l i k e l y t h a t t h e l i c k s do n o t c o n t a i n 115 enough selenium to be e f f e c t i v e i n reducing the myopathy. Since animals entering and leaving the l i c k exhibit gross symptoms of par a l y s i s , i t i s not l i k e l y that the l i c k s are supplying s u f f i c i e n t selenium. Therefore i t i s u n l i k e l y that selenium i s the at t r a c t i n g element. Rosenfeld and Beath (1964) suggest that the incidence of the disease i s higher i n areas where gypsum f e r t i l i z a t i o n i s practiced i n order to increase the y i e l d of forage. It i s inte r e s t i n g to note that gypsum mines are common i n the area and i t i s not u n l i k e l y that gypsum concentrations exist i n the' P u r c e l l Mountains, i n the v i c i n i t y of goat range. 116 Table 19 Selenium values for plant samples and earth l i c k samples from the P u r c e l l Mountains, on a dry matter basis o Description of Material Selenium i n p0pom0 Idaho Fescue o032 Thlckspike Wheatgrass 0CV3 Bluebunch Wheatgrass cOl^ Buffalo Berry Kentucky Bluegrass o032 Northwestern Sedge .0^3 Rush (Juncus parryl) o031 Alpine Strawberry ,037 Sedge Species , o029 Spike Trisetum .016 Alpine Bluegrass .079 Bear Berry o030 Alpine F i r „029 Toby Creek l i c k sample „090 Dutch Creek l i c k sample ,038 DISCUSSION AND CONCLUSIONS On examining the use of natural l i c k s by goats, i t appeared that patterns of movement place the animals, on a yearly basis, on ranges of d i f f e r i n g chemical composition,, The use of l i c k s by goats i n the summer i s an attempt to supplement th e i r diet i n r e l a t i o n to the chemical nature of the ranges. The sodium content of the forage c o l l e c t e d at a varie t y of elevations i s so low that i t does not appear to meet the needs of the animal. In the East Kootenay, almost no information has been c o l l e c t e d regarding the winter and summer ranges of the goat. The amount of time spent on any one range or the range they leave to move to natural l i c k s i s v i r t u a l l y unknown. This does not allow an estimate of the effects of a winter dietary deficiency. It appears then that the evidence presented i n t h i s thesis supports the idea that the craving is. induced i n the spring rather than over the winter. Observations of other ungulates wintering i n the v i c i n i t y of the l i c k s suggest that l i c k use occurs after feeding on spring forage high i n water and low i n sodium. The fact that goats do not move d i r e c t l y into the l i c k s upon leaving the winter range supports t h i s Idea, At Lazy Lake a feeding period of one to two weeks preceded l i c k use. Goats entering the v i c i n i t y of the l i c k s i n early spring descend to low elevations to take advantage of succulent forage i n region 2, Where thi s region i s a north facing slope or at a s l i g h t l y higher elevation goat a c t i v i t y i s delayed, as i s the growth of forage. The importance of the c r i t i c a l period i n the spring i s obvious and requires a more detailed examination from the standpoint of sodium content of urine, blood and forage along with water content of the forage. As explained, females do not move to the l i c k s u n t i l after the kidding period. It i s not known whether they u t i l i z e succulent forage prior to kidding since their winter range and kidding area i s not known. It does not seem l i k e l y that females are t r a v e l l i n g extremely long distances immed-i a t e l y after giving b i r t h . If the use of succulent forage i s as important as suggested, females would be the most l i k e l y group to take advantage of i t . While using region 2 females with kids ranged at progressively higher elevations, thereby taking advantage of succulent forage for at least a month. The r e l a t i o n of kidding, succulent forage and l i c k use Is an Interesting one that deserves attention i n the future. The population estimate given In t h i s study i s much higher than most l o c a l reports suggest, An attempt to confirm, by a e r i a l census, the estimate of the number of goats calculated to inhabit the area between the Dutch Creek and 119 Toby Creek l i c k s was unsuccessful, probably because we were unable to include the head water areas of the two drainages. It i s not known whether these animals move from the head waters of the two drainages to the Toby and Dutch Creek l i c k s i n the spring or i f they move from a closer winter range. It appears that they move to t h i s alpine range on leaving the l i c k s i n the f a l l . The winter range would appear to be the key factor i n determining movement to the l i c k s , since animals move to the l i c k s immediately after the snow recedes. Patterns of movement have been examined i n r e l a t i o n to populations, groups and i n d i v i d u a l s . Marking at the Toby Creek l i c k has shown that frequent t r i p s are made between region 2 and 3 . At both the Toby and Dutch Creek l i c k s f e -males t r a v e l longer distances to the l i c k s than do males, and make the t r i p less frequently. Females used the Dutch Creek l i c k only during peaks i n temperature and had to t r a v e l f i v e to ten times farther than did the females using the Toby Creek l i c k . At Toby Creek the c o r r e l a t i o n between female use and temperature i s very poor. It Is not known at t h i s time how important distance i s i n regulating movement between storms, Male use was correlated with temperature and storms i n May at the Toby Creek l i c k although the distance between region 2 and 3 i s very short. The storms at thi s time are severe and may overbalance the short distance. U n t i l the 120 exact boundaries of ranges can be delineated, the importance of distance i n regulating movement between storms and f r e -quency of use can not be determined,, During the summer, goats select to a very high degree, the t r a i l s they use moving to and from the l i c k s , the actual l i c k they use and within a l i c k , c e r t a i n s i t e s are preferred over others, It can not be overemphasized that r goats very seldom wander through the timber and that they use the t r a i l s between region 1, 2 and 3 to maximize their e f f i c i e n c y of movement. Due to the long distances they must often t r a v e l to reach a l i c k and the v a r i e t y of t e r r a i n they must traverse, i t can be seen that t r a i l systems are advan-tageous for their s u r v i v a l . In t h i s regard, region 2 i s very important. Upon reaching the v i c i n i t y of a l i c k , region 2 provides the animal with food and escape t e r r a i n and serves as a base from which the animal can u t i l i z e the l i c k or escape the effects of a storm. Goats appear to be able to t e l l the difference be-tween l i c k s and w i l l select the one having the highest sodium concentration. In three instances the secondary l i c k was more accessible but was bypassed i n preference for the primary l i c k . Within a l i c k , selectiveness for s p e c i f i c s i t e s appears to be quite high and at three l i c k s examined sodium was the only element consistently higher In preferred s i t e s . 121 It has been shown that sodium i s de f i c i e n t i n the forage available to goats at three elevations and at two seasons of the year. Evidence suggests that t h i s element i s available to the animals i n s u f f i c i e n t quantities In the l i c k s studied, to act as a supplement. Due to lack of data regarding sodium concentrations i n the urine, serum sodium levels could not be used to indicate a deficiency i n the animal. Since studies on other animals suggest that serum sodium can vary independently from the craving exhibited, It i s quite possible that t h i s i s the case with the goat. It appeared that white muscle disease may be present i n c e r t a i n populations of goats i n the area. According to Muth and Allaway (1963) t h i s i s a possible selenium d e f i c i e n t area. While i n the v i c i n i t y of the l i c k goats eat buffalo berry, although i t i s known that very few other ungulates w i l l do so. Since t h i s forage contained rather high amounts of selenium the sel e c t i v e feeding behaviour of the goat may be an attempt to reduce t h i s d e f i c i e n c y . The goat winters at high elevations where th i s species of forage i s r e l a t i v e l y scarce and therefore unavailable for much of the year. Feeding on vegetation low i n selenium has been known to d i r e c t l y cause white muscle disease. 122 LICKS AS A MANAGEMENT TOOL One of the f i r s t uses of natural l i c k s as a manage-ment t o o l should be to supply ungulates with desired nutrient material that i s known to be d e f i c i e n t In the area. This should be accomplished without increasing the attractiveness of the l i c k or i t s concentrating power. In the study area, selenium i s thought to be d e f i c i e n t . It seems probable that t h i s element could be supplied to the animals as sodium selenate, by placing small quantities i n high l i c k i n g s i t e s so that i t would be ingested while the animal i s eating the clay s o i l . S i m i l a r l y , phos-phorus, which becomes d e f i c i e n t i n the vegetation during the winter, could be impregnated into l i c k s o i l s for those species wintering i n the v i c i n i t y of natural l i c k s . SALTING Whether or not to include a s a l t i n g program as a part of management i s d i f f i c u l t to answer. It has not been c l e a r l y demonstrated that the absence of supplementary sodium chloride has i l l e f f e c t s on animals any more than i t has been proven that i t s presence i s b e n e f i c i a l . Altmann (1952) found that a r t i f i c i a l l y supplied s a l t does not influence elk migration to any great extent i n free ranging elk. Natural l i c k s are often determinants of elk 123 movements i n l o c a l areas. The most promising employment of s a l t i n g , then, seems to be the f u l f i l l m e n t of phys i o l o g i c a l needs and as a technique for securing proper d i s t r i b u t i o n of range use. Literature available indicates that s i g n i f i c a n t r e d i s t r i b u t i o n of big game through the use of s a l t has been attained only i n iso l a t e d s i t u a t i o n s , and then only by very c a r e f u l l y executed programs. In Oregon, i t i s stated that after three years of experimental work, sa l t has not been proven to be necessary or p a r t i c u l a r l y encouraging as a t o o l .in deer management. The introduction of s a l t , i n the case of the bighorn may actually concentrate range use rather than d i s t r i b u t e i t (Smith, 19 5*+ >. Salt placed above or below the range of greatest elk a c t i v i t y on the winter range was l i t t l e used. Such s a l t , even when v i s i b l e was not a t t r a c t i v e enough to cause many elk to enlarge their range to include i t , (Beeman, 1957). Since use of succulent spring vegetation occurs prior to l i c k use, i t seems reasonable that any revegetation program could cause r e d i s t r i b u t i o n and co n t r o l mobility i n big game by reseeding succulent vegetation varying distances from natural l i c k s . A r t i f i c i a l l i c k s placed at higher ele-vations outside the normal winter range would appear at t r a c t i v e since succulent vegetation usually appears on the winter ranges 12*f f i r s t . In the study area, a r t i f i c i a l l i c k s could be i n i t i a t e d i n region 2 adjacent to w e l l used t r a i l s . This would be p a r t i c u l a r l y b e n e f i c i a l to females which are vulnerable to early hunting seasons. In the f a l l , females are feeding on succulent vegetation at high elevations and t r a v e l l i n g to l i c k s at low elevations. Licks established i n region 2 would shorten the t r a v e l l i n g distance and keep the animals closer to escape t e r r a i n . Not to be overlooked are the r e c r e a t i o n a l values of l i c k s for photographers and n a t u r a l i s t s . Licks In wilder-ness areas may prove b e n e f i c i a l to the w i l d l i f e b i o l o g i s t i n c o n t r o l l i n g cropping, to the animal as a dietary supplement and to the general public as a source of recreation. 125 SUMMARY 1. The use of natural s a l t l i c k s by the mountain goat was examined from May, 1965 to September, I966 i n the East Kootenay region of B.C. The patterns of move-ment r e l a t i n g to l i c k use along with a chemical analysis of the vegetation, l i c k s o i l s and blood serum for sodium, constituted an attempt to show that sodium was the a t t r a c t i n g element, 2. The s e l e c t i o n of sodium as the important element was based on mineral c a f e t e r i a s , preference tests and l o g i c a l elimination of other elements c a r r i e d out by other workers. 3 . F i e l d methods, including those of trapping and marking were used to obtain serum samples and to follow movements of marked animals. Vegetation was c o l l e c t e d according to the method of Dietz et. a l . (1962) and s o i l samples were obtained from designated s i t e s after observing the l i c k i n g a c t i v i t y of goats. Laboratory procedures included those of Chapman and Pratt ( I96D for the determination of sodium and potassium i n forage, those of Stockstad (1953) and of the Association of O f f i c i a l A g r i c u l t u r a l Chemists (i960) for sodium, calcium, phosphorus and cobalt i n s o i l s . 126 Blood serum was analyzed for sodium i n the Kimberley and D i s t r i c t H ospital and the analysis for selenium was ca r r i e d out by the Chemistry Research D i v i s i o n i n Ottawa. 5. At each l i c k i n the study area animals u t i l i z e d regions 1, 2 and 3. Region 1 i s the summer range the l i c k serves, region 2 i s the base from which animals move to the l i c k and region 3 i s the l i c k i t s e l f . 6. Animals were very d e f i n i t e i n t h e i r use of t r a i l s , wandering very l i t t l e and each l i c k had i t s own t r a i l system. 7. Goats used the l i c k s from May to September de-pending on the elevation of the l i c k and during t h i s time encountered such a d d i t i o n a l r i s k s as joint use, parasites, predators and hunting pressure, 8. The seasonal pattern of movement i n the area i s such that goats move from higher to low elevations i n the spring while other ungulate species are moving from low winter ranges to higher summer ranges, 9. As the snow receded and the vegetation became green, l i c k s at low elevations or those on south facing slopes were u t i l i z e d . The use of l i c k s at higher e l e -vations began later i n the spring. 127 10. Use of l i c k s by males i n the spring was preceded by a period of feeding on succulent vegetation. This suggested that i n i t i a t i o n of use was due mainly to a high water uptake on a low sodium diet rather than a winter dietary deficiency, 11. As goats used the l i c k s during the summer they fed at progressively higher elevations i n region 2 and therefore had longer distances to move to the l i c k . 12. Within th i s period of seasonal use, male use of l i c k s preceded female use d i f f e r e n t i a l l y i n time. It was thought that female use was delayed due to kidding. 13. D i f f e r e n t i a l use occurred to some degree at four l i c k s , however there was a difference between high and low e l e v a t i o n a l l i c k s . l * f . Grouping was consistent with the behaviour of the animals as male groups were smaller than female groups. 15. T o t a l group size and es p e c i a l l y female group size corresponded to group size on summer range as given by Brandborg (1955)« 16. Observations on females i n the l i c k showed the following dominance hierarchys females with kids, females without kids and yea r l i n g s . On one occasion females showed dominance over males but encounters were too few to draw conclusions. 17* The pattern of molt coincided with d i f f e r e n t i a l use, with males and females without young and females with young molting in 8that order. It appeared that abrasion by vegetation greatly- affected molt. 18« Goats were most often observed i n the licks ln the afternoon while the morning and evening periods were usually spent feeding. 19. Animals were more often observed in the l i c k as the summer progressed and i t appeared that the longer distances being travelled at this time regulated the amount of licking and frequency of t r i p s . 20* Marking served as a basis to estimate the length of stay ln the vicinity of the l i c k , the frequency of visits and the Importance of region 2. 21* The movement of goats between regions 2 and 3 is largely controlled by storms and does not appear to be correlated with the temperature except when storms and low temperature occur simultaneously. 22. It appeared that approximately four changeovers Of animals occurred at the Toby Creek l i c k during the summer. This allowed us to estimate that approximately 70 animals were using the l i c k . 23. The vegetation i n the area contained very l i t t l e sodium and no apparent trend existed from spring to f a l l or among el e v a t i o n a l regions, 2hc Analysis of the l i c k s o i l s suggested that pre-ferred l i c k s had a higher sodium content than did less preferred ones and within a l i c k , high l i c k i n g s i t e s had a higher sodium content than did low l i c k i n g s i t e s . Calcium, cobalt and phosphorus did not exhibit a s i m i l a r trend. 25. Blood serum, analyzed for sodium, did not re-v e a l a deficiency i n the animal. At the supposed levels of deficiency serum sodium was not u s e f u l In determining sodium as the a t t r a c t i n g element since i t was too highly regulated. 26. Animals which died during trapping operations exhibited gross symptoms attributed to white muscle disease. Analysis of the vegetation and l i c k s o i l s suggested that selenium was present i n amounts causing the myopathy,, 27. Licks may serve as a useful, management t o o l . Elements d e f i c i e n t i n the area s could be placed In high l i c k i n g s i t e s so that animals could supplement their natural d i e t . 130 LITERATURE CITED ALTMANN, M. 1952, S o c i a l behaviour of elk (Cervus canadensis nelsdhi) in the Jackson Hole area of Wyoming, Behaviour 4 (2)s 116-143, ANDERSEN, N.A. 1940. Mountain goat study, B i o l . B u l l . No. 2. Washington State Dept. of Game. 21 pp. ASSOCIATION OF OFFICIAL AGRICULTURAL CHEMISTS, i 9 6 0 . O f f i c i a l methods of analysis of the AOAC. 9th e d i t i o n (Wash. D.C), BEEMAN, R.D. 1957, Salt i n the management of elk and other w i l d l i f e i n the Lower Selway River area. M.S. Thesis (unpubl.). University of Idaho, Moscow, 113 PPo BEILHARZ, S. 1962. The ef f e c t of concurrent deficiency of water and sodium on the sodium appetite of sheep. Jour, of Physiol. 163§ 378-390. BISSEL, H.D. 1953. Free choice mineral experiment. C a l i f o r -nia P.R. Proj. W-25-R, PoR, Quarterly Prog, Rept. 13 (2)? 237-BLAIR, R.M. and E,A. EPPS. I967. D i s t r i b u t i o n of protein and phosphorus i n spring growth of rusty blackhaw. J. of W i l d l i f e Mgt. 31 188-189. BRANDBORG, S.M. 1955. L i f e history and management of the mountain goat in Idaho. Wildlife Bull, No, 2. Dept, of Fi s h and Game, Boise, Idaho. 142 pp, BROWMAN, L.G. and H.S. SEARS. 1955. Erythrocyte values and alimentary canal pH values in mule deer. J, of Mamm, 36 % 474-476. CHAPMAN, H.D. and P.F, Pratt. I 9 6 I . Methods of analysis for s o i l s , plants and water. U. of C a l i f . Div. of Agric. Science, pp. 60-61. CLOVER, M.R., 1954. • Deer marking devices. C a l i f . Fish and Game. 40s 175-181. 131 COLLANDER, R. 19^1. Selective absorption of ions by higher plants. Plant Physiol, 16§ 691-720, COOK, CW. and L.E. HARRIS. 1950. The n u t r i t i v e value of range forage as affected by vegetation type, s i t e and stage of maturity. Utah Agric. Expt. Sta. B u l l . 3*Hf. ^5 PPo COWAN, I. MoT. 19^ +0. D i s t r i b u t i o n and v a r i a t i o n i n the native sheep of North America. Am.Midland N a t u r a l i s t . 2h% 505-580. COWAN, I. McT. 1 9 ^ , Report of w i l d l i f e studies i n Jasper, Banff, and Yoho National Parks and parasites, diseases and i n j u r i e s of game animals i n the Rocky Mountain National Parks of Canada. W i l d l i f e Service, Dept. of Mines and Resources, Ottawa. 83 PPo (mimeo). COWAN, I. McT. and V.C. BRINK. 19^9, Natural game l i c k s i n the Rocky Mountain National Parks of Canada. J. of Mamm. 30 (*+)§ 379-387. COWAN, I. McT. 1951o The diseases and parasites of big game mammals of Western Canada. Rept, of Proc. of the 5th Annual Game Convention. B.C. Fi s h and Game Dept. pp. 37-6H-0 COWAN, I. McT. and C J „ GUIGUET. 1965. The mammals of B r i t i s h Columbia. B.C. Prov. Museum Handbook. No. 11. hlh pp. DALKE, P., R„D, BEEMAN, F.J. KINDEL, R 0 J 0 ROBEL and T„R. WILLIAMS. I965. Use of s a l t by elk i n Idaho. J. of W i l d l i f e Mgt. 29 (2)g 319-332. DALKE, P., R.D.* BEEMAN, F.J» KINDEL, R.J. ROBEL and T.R. WILLIAMS. 1965. Seasonal movements of elk on the Selway River drainage, Idaho. J . of W i l d l i f e Mgt. 29 (2)s 333-338. DEMARCHI, R.A. 1965. An ec o l o g i c a l study of the Ashnola Bighorn winter ranges, M,Sc„ Thesis (unpubl*). Dept. of Plant S c i . University of B.C.. 103 pp. X>2 DENTON? D„A. 19640 The behaviour of sodium d e f i c i e n t sheep. Behaviour, 20g 364-376. DENTON5 D.A. and J.A. SABINE. I96I. The sel e c t i v e appetite for sodium shown by sodium d e f i c i e n t sheep. J. of Physiol. 157s 97-116. DEPARTMENT OF TRANSPORT. 1965. Climate-logical reports for the Cranbrook a i r p o r t . V i c t o r i a , B.C. DEPARTMENT OF TRANSPORT. I966. C l i m a t o l o g l c a l Reports for the Cranbrook airport and Radium Junction. V i c t o r i a DEVLIN, T.J. and W.K. ROBERTS. 1963. Dietary maintenance requirements of sodium for wether lambs. J. of Animal S c i . 22a 648-653-DIETZ, D.R*, R.H. UDALL and L.E. YEAGER«, 1962, Chemical com-position and d i g e s t i b i l i t y by mule deer of selected forage species, Cache l a Poudre Range, Colorado. Tech. B u l l . No, 14, Colorado Game and F i s h Dept. 89 pp. DUKES, H*H. 1955. The physiology of domestic animals. Comstock Publishing Associates, Comstock, New York, 1020 pp. ETKIN, W„ 1964, S o c i a l behaviour and organization among vertebrates. The University of Chicago Press, pp, l-3^o FORBES, G,B, I962. In C.L. Comar and F, Bronner (ed.). Mineral metabolism. Academic Press, New York, pp. 2-72. FOREST SERVICE. 1937c Range plant handbook. U..S. Dept. of Agric. U.S. Gov't. Printing O f f i c e . Washington, D.C. GEIST, V. 1964. On the r u t t i n g behaviour of the mountain goat. J. of Mamm. 45 (4)g 551-568, GILBERT, F„A, 1957, Mineral n u t r i t i o n and the balance of l i f e . University of Oklahoma Press, 350 pp. GORDON? J.G., D 0E. TRIBE and T.C GRAHAM.. 195*+. The feeding behaviour of phosphorus-deficient c a t t l e and sheep. Nut. Abs. 195S 24- (No. 53^7)« 9^8. HANSEN j C.G, 1963. A dye spraying device for marking desert bighorn sheep. Desert Bighorn Council. 7? 199-202. HARMER, P'.M. and E*J. BENNE. 195+5„ Sodium as a crop nutrient. S o i l S c i . 60s 137°l*+8. HIDIROGLOU, Mo 1965. Influence of selenium on the selenium contents of hair and on the Incidence of n u t r i t i o n a l muscular disease i n c a t t l e . Can. J. of Animal S c i , H-58 197-202, HITCHCOCK, A.S. 1950. Manual of the grasses of the U.S. 1950. Misc. Publ. No. 200, Dept. of Agric, Washington, D.C, 10^0 pp, HOLLAND, S.S, 196*+. Landforms of B r i t i s h Columbia. A physiographic o u t l i n e . B u l l . No. *+80 B r i t i s h Columbia Dept. of Mines and Petroleum Resources. 138 pp. HONESS, R,F. and N . M , FROST. 19^2. A Wyoming bighorn sheep study. B u l l . No. 1. Wyoming Game and Fish Dept. 127 PP . HUFFAKER, R„C. and A. WALLACE. 1959. Sodium absorption by di f f e r e n t plant species at d i f f e r e n t potassium l e v e l s . S o i l S c i . 8?§ 130-13^. KELLEY, C C . and W.D. HOLLAND, I96I . S o i l survey of the Upper : Columbia River V a l l e y i n the East Kootenay D i s t r i c t of B.C. Rept. No. 7 of the B.C. S o i l Survey. B.C. Dept. of Agriculture. 107 pp. KERR, D. I965. Mountain Goat Ecology i n Alberta. M.Sc. Thesis (unpubl.). University of Alberta, Edmonton, KITTS, W . D . , P . J o BANDY, A . J . WOOD and I„ McT. COWAN. 1956. E f f e c t of age and plane o f n u t r i t i o n on the blood chemistry of the Columbian black-t a i l e d deer (Qdocoileus hemionus columbianus). Can. J . o f Zool. 31+ % if77 . i4 .g i f . KLEIN, D.R. 1 9 5 3 . A. reconnaissance study of the mountain goat i n Alaskao M.Sc0 Thesis (unpubl.)» University of Alaska. College , KRAJINA, V . J o 1 9 6 5 . Ecology of Western North America. Dept. of Botany, University of B.C. 112 pp. LEHR, J . J . 1 9 5 + 1 „ The importance of sodium for plant n u t r i t i o n . S o i l . S c i . 52s 237-2M+, LYONS, C P . 1 9 6 5 . Trees, shrubs, and flowers to know In B.C. Dent and Sons (Canada) Limited. 19*+ pp. MARRIOT? H.L. 1950. Water and s a l t depletion. Ryerson Press, Toronto. 80 pp. McLEAN, A. and E.W. TISDALE. i 9 6 0 . Chemical composition of native forage plants i n B.C. i n r e l a t i o n to grazing practices*, Can. J , of Plant S c i . h-Os ^ 0 5 - ^ 2 3 . McLEAN, A,, H.H. NICHOLSON and A.L. VAN RYSWYK. 1963. Growth, productivity and chemical composition of a sub-alpine meadow i n Interior B DC 0 J . of Range Mgt. 1 6 ( 5 ) * 2 3 5-2^ 0 . MUTH, O.H. 1 9 6 3 . White muscle disease, a selenium-responsive myopathy. J . of Amer. Vet. Med. Assoc. IH-2 (12) S 1379-138^. MUTH,'O.H. and W.H. ALLAWAY. I963. The r e l a t i o n s h i p of white muscle disease to the d i s t r i b u t i o n of naturally occurring selenium. J . of Amer. Vet. Med. Assoc. l*+2 (12)s 1379-138^. PACKARD, P.M. I9V60 An e c o l o g i c a l study of the. bighorn sheep i n the Rocky Mountain National Park, Colorado,, J . of Mamm. 27 C D s 3 - 2 8 . PAYNE, J.M. 196k0 Factors a f f e c t i n g plasma calcium and i n -organic phosphorus concentration In the cow with particular reference' to pregnancy, l a c t a t i o n and age. Nut. Abs„ 19659 35 (No. 573)s 95« ROSEN, M.N. and A.I. BISCHOFF. 1952. The r e l a t i o n of hematology to condition i n C a l i f o r n i a deer. Tran. N.A. W i l d l . Conf. 17s ^ 8 2 - ^ 9 6 . ROSENFELD, I„ and O.A. BEATH. 1964 . Selenium. Academic Press , N.Y,. 411 pp. SAUNDERS, J.K. 1 9 5 5 . Food habits and range use of the Rocky Mt. goat i n the Crazy Mountains, Montana. J. of W i l d l . Mgt„ 19 (4)s 4 2 9 - ^ 3 7 . SELLERS, A.F. and M.H. ROEPKE. 1951 . Studies of ele c t r o l y t e s i n body f l u i d s of dairy c a t t l e . 1) E f f e c t s of water diuresis on renal excretion of several elements. Am. J. of Vet. Research 1 2 . 183-186 , SMITH, S.E. and P,D, AINES. 1959» Salt requirements of dairy cows. C o r n e l l Agric. Expt. Sta. B u l l . 9 3 8 . 26 pp. SMITH, D.R. 1 9 5 4 . The bighorn sheep i n Idaho, i t s status, l i f e h i s t o r y , and management. W i l d l . B u l l , No. 1. Idaho Dept. of Fish and Game. 154 pp, STOCKSTAD, D.S. 1 9 5 3 , Chemical c h a r a c t e r i s t i c s of natural l i c k s used by some big game animals i n western Montana. Trans. N.A. W i l d l . Conf. 1 8 : 247-2 5 8 . STUFFLEBEAM, C E . 1964 . Seasonal variations i n levels of some chemical and haematological components i n the blood of Hereford cows. Nut. Abs. 1 9 6 5 5 35 (No. 3 9 1 D * 6 8 3 . VRZGULA, B. 1 9 6 2 . Sodium, potassium and calcium'In blood serum of young c a t t l e i n r e l a t i o n to season. Nut. Abs. 1 9 6 3 , 33 (No. 6 1 9 3)2 1 0 2 0 . VRZGULA, B. I 9 6 5 . Influence of age i n n u t r i t i o n , kalium, and calcium levels of Bovine blood serum. Nut, Abs. 1 9 6 5 , 35 (No. 2240)s 3 8 8 . WALLACE, A., S.J. TOTH and F . E . BEAR. I947. Sodium content of some New Jersey plants. S o i l S c i . 65» 249=258. WE IS BERG, H.Fc 1953 c Water, e l e c t r o l y t e and acid-base balance. The Williams and Wilkins Co., Baltimore. 245 pp. " WILLIAMS, T.Rc 1 9 6 2 . The significance of s a l t and natural l i c k s In elk management. M.Sc, Thesis (unpubl.)o University of Idaho, Moscow. 200 pp. 136 APPENDIX Ao SCIENTIFIC AND COMMON NAMES FOR PLANT SPECIES MENTIONED References includes Hitchcock Forest Service.(1937). (1950), Lyons (1956), and 137 Grasses and sedgess Aeropvron dasystachyum thickspike wheatgrass Aeronvron spicaturn bluebunch wheatgrass Calama^rostis rubescens pine grass Carex concinnoides northwestern sedge Car ex s.p.p, sedge Festuca idahoensis Idaho fescue Koelerla c r i s t a t a June grass Poa alplna alpine bluegrass Poa pratensis Kentucky bluegrass Stipa columblana subalpine needlegrass Stipa r i c h a r d s o n i i Richardson needlegrass Trisetum spicaturn spike trisetum Forbs: Fraearia glauca strawberry Shrubs and trees s Abies lasiocarpa alpine f i r Atnelanchier aInifolia Saskatoon berry Arctostaphylos uva°ursi bear berry Juniper us scopular-um Rocky Mountain juniper Populus tremuloides trembling aspen Pseudotsuga menziesii Douglas f i r S a l i x spp. willow Shepherdia canadensis buffalo berry Vaccinium scoparium grouse berry 

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