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Home My WebLink AboutAPA911ALASKA DEPARTMENT OF FISH AND GAME J U N E A U, A L A S K A STATE OF ALASKA William A. Egan, Governor DEPARTMENT OF FISH AND GAME Wallace H. Noerenberg, Commissioner DIVISION OF GAME Frank'Jones, Acting Director Donald McKnight, Research Chief MOOSE RESEARCH REPORT by Robert E. LeResche and James L • Davis Volume XII Project Progress Report Federal Aid in Wildlife Restoration Project W-17-3, Jobs l.lR, 1.2R, 1.3R, 1.4R Persons are free to use material in these reports for educational or informational purposes. However, since most repo~ts treat only part of continuing studies, persons intending to use this material in scien- tific publications should obtain prior permission from the Department of Fish and Game. In all cases tentative conclusions should be identified as such in quotation, and due credit would be appreciated. (Printed September, 1971) :~t~~;L4 ( ACKNOWLEDGMENTS The following Alaska Department of Fish and Game and United States Fish and Wildlife Service personnel participated in scientific activities relevant to this report: D. Bader, J. Coady, D. Cornelius, J. Didrickson, s. Eide, C. Erskine, L. Glenn, P. Havens, J. Hemming, c. Jackson, A. Johnson, L. Johnson, R. Kramer, J. Kurtz, P. LeRoux, C. Lucier, L. Miller, J. Oldemeyer, R. Pegau, R. Perkins, K. Pitcher, J. Reynolds, J. Sexton, R. Somerville, N. Steen. R. A. Rausch was instrumental in implementing all the work reported here and offered many helpful suggestions. C. Mcilroy developed the technique for counting rumen protozoans, aged moose teeth, and gave advice and assistance on other laboratory projects. J. Hakala, R. A. Richey and R. K. Seemel cooperated extensively with the project on logistic and scientific levels and provided some unpublished data cited here. G. Watson, U. S. Bureau of Sport Fisheries and Wildlife, Anchorage Region, kindly allowed use of a computer for data analyses. We sincerely thank all the above, without whose assistance this report would not be possible. State: Cooperators: Project No.: Job No.: JOB PROGRESS REPORT (RESEARCH) Alaska Alaska Department of Fish and Game, U. S. Bureau of Sport Fisheries and Wildlife, Kenai NWR W-17-3 l.lR Project Title: Moose Investigations Job Title: Moose Productivity & Physiology Period Covered: July 1, 1970 through June 30, 1971 SUMMARY Natality, mortality and yearling recruitment of moose (Alces alces gigas) were determined in four one-square-mile pens. During the period June 1968 through June 1971, calf production ranged from 43 to 72 calves per 100 adult females (mean 58:100). Yearling recruitment varied between 11 and 41-45 yearlings per 100 adult females (mean 25-27). Adult mortality almost balanced yearling recruitment, with an overall change over four years of + 3 percent of the adult population. · Mortalities exceeded proportion of presence in the penned populations among calves (28 percent of recorded mortalities vs. 21 percent presence in the population P) .20; t-test) and animals 12-24 months of age (24 per- cent vs, 14 percent P ( .10; t-test). Mortalities of females over 24 months of age (48 percent of mortalities) were fewer than adult females' propor- tion in the population (55 percent) (P ) • 20, t-test), On 1 July 1971 the pens contained 29 adult females, 22 calves, 13 yearlings and 11 adult males: total 75 moose, or a mean density of 18.8 moose per square mile. Age structures of penned populations were equiva- lent to one another and to the population immediately outside the pens (P ) .10; x2 test), Since July 1969, 417 sera, 174 whole blood specimens and 170 smears were collected from moose of all ages during all months of the year. Sera were analyzed by standard auto-screening (SMA-12) processes and electro- phoresis for 16 parameters. Whole blood specimens and smears were analyzed for standard hematologic values. Multiple regression analyses are being performed on all parameters with respect to season, location and method of collection and age, sex, reproductive condition, body measurements and body weight of the moose. Calculations are not complete. Sera from 12 winter-killed moose showed depressed levels of calcium, phosphorus, glucose, uric acid, total protein, albumin and a depressed albumin/globulin ratio. BUN was elevated 250 percent over previously estimated "normal" values. i Twig-count methods for determining browse utilization proved more precise in heavily-overused range than in normally browsed range. Forty whole body weights of moose were obtained and a weight/age curve was constructed. Body weight fluctuated seasonally as much as 30 percent over basal June weight, but June and September weights appeared to increase progressively through at least the 13th year of life. Cows with calves weighed 8-18 percent less in summer than the same or similar individuals when without calves. Girth/total length ratio was found to be a probably valid indication of relative weights of sequentially-handled individuals, but was not valid as a weight estimator for different individuals. Food intake and activity patterns of three tame moose (two males) were observed for 99 hours during summer on normal range, for 19 hours in winter on normal range and for 30 hours during winter on depleted range. Moose consumed approximately the same number of bites of forage each day in winter and summer, but the consumption was concentrated into approxi- mately two-thirds the time in winter. In winter, moose spent a greater proportion of each daylight hour in feeding (61 percent vs. 43 percent in summer) and consumed slightly more bites per hour of feeding (730 vs. 646). Food eaten varied between summer and winter, and moose ate a greater variety of forage during all seasons than previously realized. Birch (Betula papyrifera) leaves comprised 56 percent (by number of bites) of the summer diet, forbs 25 percent, grasses, sedges and aquatics 10 percent and willow (Salix ~) .5 percent. In winter, diet on range that had supported average moose numbers for the area was 72 percent birch twigs, 21 percent lowbush cranberry (Vaccinium vitis-ideae), and 6 percent willow and alder (Alnus crispa). On depleted winter range, stocked for 18 months with abnormally high moose densities, birch twigs composed only 22 percent of the diet. The bulk of bites taken were of lowbush cranberry (51 per- cent) and fruticose lichens (23 percent). Bites cranberry: bites birch and bites lichen:bites birch ratios reached 10-30 in May. Identifiable food from moose killed in early December on climax willow range was (by volume) 59 percent willow, 33 percent birch and 8 percent aspen (Populus tremuloides). Similar values for animals in sera! birch range were 39 percent willow, 36 percent birch, 14 percent lowbush cranberry and 6 percent aspen. Willow was selected out of proportion to its abundance in the sera! range. Numbers of rumen protozoa per cc. of rumen liquor and proportion of microorganisms (by volume) in the liquor were greater on climax willow range than on sera! birch range, indicating perhaps the former range type is of higher quality. Snow cover at the Moose Research Center was a minor factor in limit- ing available food for moose during 1970-71. ii SUMMARY •. BACKGROUND OBJECTIVES PROCEDURES FINDINGS . CONTENTS Productivity and Mortality Within Pens ... Blood Values as Indicators of Nutritional Status .••.. Browse Production and Utilization • Weights and Measurements. • • • •. • • Tame Moose Feeding Habits ••••.• Food Habits from Rumen Analysis . Chemical Composition of Forage Plants . Snow Studies .••••••• Miscellaneous Observations. RECOMMENDATIONS .•••.. LITERATURE CITED . BACKGROUND Page No. i 1 7 7 14 14 • 38 . 41 . 43 47 • 61 70 • 70 70 70 73 Management of moose (Alces alces) in Alaska has always relied on gross knowledge: 1) of population indices (eg: relative numbers, sex ratios, age composition, apparent natality) gained through observation (usually by air) of large numbers of animals; 2) of hunter harvest learned from reporting procedures; 3) of unusual mortalities ("die-offs") when noticed; 4) of in utero pregnancy rates; and 5) of occasional crude and subjective observations of browse conditions. The state's expanding human population makes these methods, by themselves, obsolescent for various biological and political reasons. For example, on the Kenai Peninsula, an extensive (more than 350,000 acres) burn occurred in 1947, with a subsequent striking increase in the number of moose present (Spencer and Hakala, 1964). By the late 1950's or early 1960's, numbers seemed to have stabilized at a high level; yet the predominately birch seral type was noticably browsed in very few areas, and hedged in even fewer places. Moose production had fallen behind browse production. No major "die-off" had been recorded, browse was not (to the superficial observer) over-used, hunter harvest was low in relation to the moose population size, and most (more than 90 percent) cow moose were pregnant in fall and early winter. Less-than-obvious factors were at work. More sophisticated knowledge of and means of quantifying moose physiological and vegetation parameters were needed. Situdtions requiring such knowledge are not localized on the Kenai Peninsula. The information is needed for solving present and future 1 problems wherever moose occur in the state. The Tanana River flats near Fairbanks support a moose population similar in density to that in the Kenai 1947 burn. In spring 1965, gross signs of loss of condition were observed, few viable calves were produced, and collections revealed severely undernourished calves. Classical symptoms of severely browsed shrubs and high moose densities were present, but more specific analysis was impossible given the then-and-present state of knowledge. At about the same time (1962 through 1965) moose numbers apparently declined markedly in the Nelchina Basin: no explanation but severe winters was possible. Matanuska Valley populations, the only ones in the state even possibly manipulated by hunting, fluctuate in ways that could be better understood and predicted by more specific knowledge of moose-habitat interrelationships than is currently available. During the winter and spring of 1970-71, die-offs occurred in at least two areas of the state: the Tanana Flats and various drainages on the west side of upper Cook Inlet. These winter snow-caused mortalities will likely affect moose numbers and browse for at least several years. The overall project objective of jobs carried out at the Kenai Moose Research Center is to obtain a more thorough and specific knowledge of how moose affect vegetation and how vegetation affects moose. Within the last decade, classical theories of ungulate population regulation have been increasingly questioned. The idea of moose popula- tions being controlled catastrophically by quantity of available winter range (eg: Spencer and Chatelain, 1953), while still the best explanation of some population_s' fluctuations, did not fully describe others. As more data became available, nuances of selective mortality by age and sex and the importance of range quality (Klein, 1970; Flook, 1970) to Cervids became more evident. Some populations appeared to be self-regulating without catastrophy. These populations were often characterized by low productivity and medium to high densities (Wynne-Edwards, 1970; Cole, 1971; Houston, 1971). The ecosystem approach to such problems has become increasingly popular as methods have developed. The approach is a multi-variant one, with feedback mechanisms, predation, food quality and quantity and other variables considered from an energy-flow standpoint (cf: Petrides et al, 1968; Lewis, 1969; Wagner, 1969). To date, probably no big game popula- tions are well enough known for study by proper ecosystem-type analyses. The one attempt to thus consider a moose population, though confined primarily to only a consideration of biomass, (Jordan and Botkin, 1970) failed because of many incomplete data being inserted into the model. The logical first step in determining energy-flow relations in an ungulate ecosystem is determination of the energetics of individual animals. Several ungulate studies to date have considered energetics of wild herbivores (eg: Abrams, 1968; Rogerson, 1968; Silver et al, 1969; Silver ~ al, 1971) but none have investigated moose. Similarly, recent studies have evaluated nutritional requirements of ungulates other than moose (eg: Nordan et al, 1968; Taylor, 1968; Ulbrey et al, 1969; McEwan and Whitehead, 1970)-.----- 2 The study of animal nutrition as it affects reproduction is especially relevant to considerations of population regulation, especially in moose, where maximum mortality probably occurs within several days postpartum (LeResche, 1968). Studies of ungulates (eg: Thomson and Thomson, 1948; Verme, 1962; Verme, 1969) and other mammals (eg: McLean and Usher, 1970; Nobmann and Adams, 1970; Shoemaker and Wurtman, 1971) confirm the impor- tance of good nutrition to maximal reproduction. Moose research dealing with nutritional aspects of productivity has been slight. Most work has involved a population approach to learning productivity, mortality and recruitment, often by aerial counts (eg: Denniston, 1956; Pimlott, 1956, Spencer and Hakala, 1964; Knowlton, 1960; Rausch and Bratlie, 1965; Simpkin, 1965; Rausch and Bishop, 1968; Bishop, 1969). Fewer studies have used physiological or behavioral approaches to learning productivity (eg: Rausch, 1959; LeResche, 1968; Houston, 1968; Markgren, 1969). No other productivity records for individual moose (except by ovarian analyses) over more than one season appear in North American literature (cf: LeResche, 1970), although Soviet domestication work at Pechora-Ilych Reservation (Knorre, 1961) has undoubtedly gathered such information. Evaluation of the nutritional status of animal populations remains difficult despite many newly suggested techniques (Jeliffe, 1966). Analyses of urine (Blaxter et al, 1966), epithelial tissue (Nutr. Res. 1970), saliva (Murphy and Connell, 1970), blood constituents and body weight and size have been attempted as indicators of nutritional status. The latter two methods were selected as most practical and promising for use on moose populations. Blood studies of moose and other wildlife species have been few and superficial and only sometimes related to nutrition. Braend (1962) con- sidered blood groups in moose, Nadler et al (1967) studied serum proteins and transferrins, and Houston (1969) analyzed several serum parameters from 13 moose. Dietrich (1970) reported hematology of six moose and several other arctic mammals. More thorough nutrition-related studies have been carried out on other Cervidae. Herin (1968) reporte~ 14 blood parameters for 39 elk (Cervus canadensis). Kitts et al (1956) related age and nutrition to hematological values in black-tailed deer (Odocoileus hemionus ~). More recently, Seal and Erickson (cf: 1969) have begun quite sophisticated blood studies of several large mammals. LeResche (1970) published serological and hematological values for more than 250 moose, but considered the data incomplete at that time. Nevertheless, he suggested that several parameters (calcium, phosphorus, BUN, uric acid, cholesterol, albumin SGOT) were promising as indicators of nutritional status. Literature regarding domestic animal and human clinical chemistry is much more complete than that concerning wild animals (cf: Swenson, 1970; Davidson and Henry, 1969 for major review texts). This literature has established the value of blood analyses in nutritional and other studies. Useful parameters include calcium (Kendall et al, 1970; Ramberg et al, 1970a; Ramberg et al, 1970b) glucose (Khan et al,-r970; Young et al 1970), urea (Preston-e~al, 1965; Eggum, 1970; Metzger et al, 1970~ ----- 3 protein (Kuttler and Marble, 1960; Beaton and McHaney, 1966), fatty acids (Mackenzie et al, 1970), insulin (Trenkle, 1970), hematologic values (Anderson et al, 1970), and others (Davidson and Henry, 1969). Special problems are presented when one attempts to evaluate nutri- tional status of wild animals using serologic and hematologic techniques. Sampling may be done only when animals are captured or killed and serial sampling of individuals is often impossible. Further, it is seldom possible to adequately evaluate food intake or environmental stresses. One of the ntost important problems in studying wild animals is that they are unaccustomed to restraint, and the very collection of samples can result in stresses that alter the parameters measured. Few studies have evaluated these factors satisfactorily. Wilson (1970) reviewed stressor agents on domestic animals and their effects, including some mention of changes in blood values. Leise et al (1970) studied effects of restraint on enzymes in leucocytes of rabbits-.-The uncertainty involved in such measurements indicates the importance of standardizing collecting methods as much as possible and of exercising extreme caution in evaluating blood constituents (eg: glucose) most reactive to the acute stresses of capture. Measurements of body weight and morphometry are potentially valuable techniques in wild animal studies primarily because data are easy to obtain and values are not subject to acute stress-related changes. Although not acutely variable, whole body weight is subject to certain variation because of changes in rumen volume (Ledger, 1968), genetic characteristics (Tamer et al, 1970) and normal seasonal variability (Verme, 1970), Vary- ing proportions of body constituents (eg: fat, muscle, water) (Short et al, 1969; Tamer et al, 1970; Seltzer et al, 1970) by season, nutrition, etc., further complicate this measure~n~of live animals. Growth patterns and body measurements, especially when used in conjunction with body weights (Klein, 1968; Wood and Cowan, 1968; Verme, 1963, 1970), minimize some of these problems if valid baseline values are available. A tremendous number of herbivore/range studies have concentrated on the range itself, with little knowledge of animal health, reproduction parameters, etc. Many of these studies have inferred animal abundance from evidence of plant use. Studies have, in the main, concentrated on browsed species, (eg: Julander, 1937; Young and Payne, 1948), but others (eg: Harlow, 1959) have taken a broader approach. Since many big game species commonly inhabit sera! ranges, many management-oriented studies have concentrated on factors affecting plant succession (cf: Heinselman, 1954; Patton and McGinnes, 1964; Halls and Epps, 1969; Mutch, 1970), Others have investigated herbivores' effects on succession and production (Garrison, 1953; Hendrick, 1958; Lay, 1965; Crouch, 1966; Krefting ~ al, 1966; Bergerud, 1968; Goodiman and Marquis, 1969; Jordan and Rushmore, 1969) and artificial ways of slowing succession or restoring a sera! state (Gibbens and Schultz, 1962; Pienaar, 1968; Leege, 1969). A major stumbling-block in range-oriented studies has been the difficulties involved in developing statistically valid browse production and utilization measuring methods (cf: USDA, 1970). These problems result in part from the extreme variances found in most random measurements 4 (LeResche, 1970), which suggest nonrandom utilization. Further difficulties stem from previous lack of exact knowledge of how many animal-days use a unit of range was sustaining. Previous studies have estimated forage pro- duction primarily by canopy cover (Evans and Jones, 1958; Goebel et al, 1958; Daubenmire, 1959; Peek, 1970), or stem counts and measurement (Bishop, 1969). Many studies have estimated browse utilization by time-consuming methods of twig counting (Heady et al, 1959; Shafer, 1963; Telfer, 1968; Bishop, 1969), but these methods~ave proven too imprecise in instances where they have been sufficiently tested (LeResche, 1970). Other methods involving browse form class are in wide use in survey-type work (Cole, 1963; Patton and Hall, 1966) but present no precise animal-use estimates. Since utilization estimates from plants are impossible with current techniques, many studies examine animals and their behavior to discover patterns of use. Most of these studies (eg: Bassett, 1951; McMillan, 1953; Harry, 1957; Knowlton, 1960; McMahon, 1964; Houston, 1968; Bell, 1970; Nicholson et al, 1970) are concerned primarily with the kinds of food eaten, but rn;ny-also attempt to estimate quantity of food consumed (cf: Van Dyne, 1968). Common techniques range from feeding trials of captive or domesticated animals (Palmer, 1944; Bilby, 1968; Reid, 1968; Marsh et al, 1971; Ulrey et al, 1971) to micro-and macro-analyses of stomac~contents or feces:from killed or living animals (Mulkorn and Anderson, 1959; Brusven and Mulkern, 1960; Storr, 1961; Bear and Hansen, 1966; Stewart, 1967; Field, 1968; Gaare, 1968; Sparks, 1968; Sparks and Malechek, 1968; Veckert, 1968; Galt et al, 1969; Hansen and Flindere, 1969; Nellis and Ross, 1969; William~ 1969; Medin, 1970; Rice, 1970; Ward, 1970), chemical methods (Theurer, 1970) and observations of wild (Harper et al, 1967; Miller, 1968; Houston, 1968) or tame (Bjugstad et al, 1970; Bergerud and Nolan, 1970; Hungerford, 1970; Laycock and Price,-r970; Martin, 1970; Nixon et al, 1970; Short, 1970; Wallmo and Neff, 1970; LeResche et al, 197l)animals. Tame animal studies solve most of the problems associated with measuring use from what is left behind and demonstrate food habits not obvious when browsed plants are observed (LeResche et al, 1971). However, other variables are introduced when tame animals are dealt with. Feeding behavior of tamed animals may be altered by taming or by supplementary feeding necessary to tame the animals. Little objective evidence is available to dispute these problems, but much empirical data (Buechner, 1950; Wallmo, 1951; McMahan, 1964; Wallmo and Neff, 1970) suggest these are minimal, especially if supplemental feeding ceases when observations begin. Problems of individual variation and quantification are more serious (cf: Wallmo and Neff, 1970). Once range production and response to utilization and species compo- sition and quantity of food consumed are known, the elements of food quality become important (Klein, 1970), Chemical composition, palatability and digestibility all determine the ultimate usefulness of a gram of available forage to a particular animal in a particular physiological state. 5 Early chemical composition studies stressed analyses of protein and sometimes phosphorus content of forages (Einarsen, 1946; Jameson, 1952; Swank, 1956; Taber, 1956; Murphy and Coates, 1966; Boyd, 1970; Klein, 1970b). These values were found, in a general way, to be directly related to range quality as indicated by ungulate browsing level, survival and reproduction. Later, proximate analyses, for crude protein, crude fat, crude fiber, ash and nitrogen-free-extract were commonly used to define forage quality (eg: Swift, 1948; Smith, 1957; Halls and Epps, 1969). Many more complex analyses of plant material (Van Soest, 1964; Short, 1966; Dietz, 1970) were undertaken, and very complete chemical and energy data are now available for many plant species (Hamilton, 1958; Cowan et al, 1970). Nutritive value of plants was found to vary with many factors (Oelberg, 1956), including site (Cook, 1959), time since burning (Cowan et al, 1950; DeWitt and Derby, 1955), season (McConnell and Garrison, 1966; Tew, 1970), palatability and digestibility. "Palatability" is a conglomerate term and includes elements of food selection, nutritive value and digestibility, as well as inherent vari- abilities in tastiness of a plant to a given animal. All studies have shown that what an animal eats is almost invariably the most nutritive and digestible forage available. Thus, discussions of "palatability" are in truth considerations of digestibility and nutritive values (eg: Albrecht, 1945; Plice, 1952; Heady, 1964; Longhurst et al, 1968, 1969). That some plants were high in nutrients and/or energy, but little eaten by herbivores led to considerations of digestibility factors and the discovery of digestive inhibitors. Regardless of its nutritive value as determined chemically, a plan~'s usefulness to its consumer is only as great as its digestibility (Johnston et al, 1968; Ulrey et al, 1970). Digestibility has been found to vary by season (Christian et al, 1970; Haggar and Ahmed, 1970), by mixture of forages consumed (Dror et al, 1970), by various plant constituents that decrease forage digestibility by their own inherent undigestibility (Short, 1966; Van Soest and Jones, 1968; Jones, 1970; Rittenhouse et al, 1970; Short and Reagar, 1970) or by inhibiting microbial action in the rumen and thereby decreasing digest- ibility for all food taken in (Oh et al, 1967; Oh et al, 1968; Mueggler, 1970). Techniques for determining~igestibility include feeding trials (Short, 1966), and in vitro (Tilley and Terry, 1963; Pearson, 1970) and in vivo (Short, 1970) methods. Concurrent with in vivo digestibility techniques, studies of rumen metabolism are of use-rn determining dietary quality of domestic and wild ruminents (Nagy, 1970). These studies range from simple field-adapted collecting and processing procedures (Klein, 1962; Short et al, 1966; Klein, 1970; Klein and Schonheyder, 1970) to comprehensive treatises on microbiology and energetics (eg: Blaxter, 1962; Hungate, 1966), Studies relevant to wild animal work have considered rumen structure (Hofmann, 1968), micro-organisms (Hobson and Mann, 1968; McBee et al, 1969; Coen and Dehority, 1970; Takayama et al, 1970) and specific food substances (Lough and Garton, 1968; Clarke and Hawke, 1970). Volatile fatty acid 6 determination has proved most useful in many practical studies (Maloiy et al, 1968; Oh et al, 1969; Dror et al, 1970). OBJECTIVES To measure natality, mortality and general condition of moose within four one-square-mile enclosures. To establish baselines by season, age and sex for the following serological and hematological parameters in moose and to evaluate their usefulness as indicators of nutritional status in moose: A. calcium B. inorganic phosphorus C. glucose D. urea nitrogen (BUN) E. uric acid F. cholesterol G. total protein H. albumin I. albumin/globulin ratio J. alpha-1, alpha-2, beta and gamma-globulins K. bilirubin L. alkaline phosphatase M. lactic dehydrogenase (LDH) N. glutamic oxalacetic transaminase (SGOT) o. hemoglobin P. hematocrit Q. white blood cells R. differential cell count (including segmenters, lymphocytes, eosinophils, monocytes, basophils) To estimate browse production and utilization and quantitatively and qualitatively estimate consumption of all plant material by moose. To learn changes in rumen protozoa levels in moose on various winter diets. To learn nutritional values and digestibilities of the more common moose forage species of plants. PROCEDURES General Description of the Moose Research Center FaciliS[ The Kenai Moose Research Center comprises four one-square-mile enclosures located in the area of the 1947 burn near Kenai, Alaska. These enclosures contain representative vegetation of both burned (regenerative: predomi- nately birch Betula papyrifera and white spruce Picea glauca) and remnant: (mixed birch-spruce-aspen Populus tremuloides stands). Marshland typical 7 of summer range is included as are well-drained hillocks supporting winter· browse-species. The entire area has been type-mapped into 11 vegetation types, and soil profiles of representative types have been completed. One hundred and forty permanent plant-succession measuring plots have been installed subjectively (five in each pen for each of seven major vegetation types), and each has been read once. A modified Daubenmire (1959) canopy estimate was employed. Eight hundred and forty permanent browse production/utiliza- tion plots have been established randomly within habitat types and these have been used to measure production (once within two of the four pens) and use (four times in one pen, three times in one pen, and once in two other pens). Twig-count and clipping methods are employed. Five five- acre exclosures are present, at least one within each enclosure. Nine fenceline traps were constructed during the reporting period, bringing the total to 21; twelve of which are within pens and nine of which are on the outside of the fenceline. Fig. 1 is a generalized map of the facility showing traps, exclosures, etc. The log headquarters building sleeps eight, and is accessible by road during dry seasons. Two-mile-long Coyote Lake provides access by float or ski plane. The Center may be reached by light plane from Anchorage in one-half hour. Populations of moose within the enclosures as of February 1, 1970, five months after enclosing pens 3 and 4, were: Pen Cows Calves Bulls Total 1 5 0 2 7 2 9 1 2 12 3 7 4 1 12 4 11 5 2 18 32 10 7 49 Pens 1 and 2 will be left unmolested in terms of moose numbers, allowing the populations to increase, decrease, or remain constant as they will. Pen 3 will be retained at its present population level and sex structure, as representative of extra-pen populations in this area, Pen 4 will be stocked with as many as 50 moose (four to five times "normal" density). Table 1 is a history of major events in construction of the facility and provides reference as to timing of events leading to the current description, 8 1 /0-1 ID-2. 10-~ CJ TRAPS 2-' 40-4 ON.E M\LE' Figure 1. Diagramatic layout of Kenai Moose Research Center. 2 2-2. I 4-2 j 3-2 t H ..3-1 4·1 40-3 Table 1. Chronology of establishing the Kenai Moose Research Center. June 1966: Construction begun. September-October 1967: January 1967: January 1968: April 1968: 1968: April 1969: June 1969-January 1970: June-July 1969: August 1969: October 1969: January-February 1970: April 1970: May 1970: August 1970: November 1970: March 1971: May-June 1971: June 1971: Browse production estimated in Pens 1 & 2. Successional plots established and read in Pens 1 & 2. Pens 1 & 2 enclosed. Moose in Pens 1 & 2 collared. Browse utilization estimated in Pens 1 & 2. Yearling bull introduced into Pen 1. Browse utilization estimated in Pens 1 & 2. Eleven traps constructed in all pens. Blood collections begun. Successional plots established and read in Pens 3 & 4. Pens 3 & 4 enclosed. TWo male calves introduced into 10-acre pen in Pen 2. Numbers of moose in Pens 3 & 4 determined. Replicate count experiments conducted. Browse utilization estimated in all four pens. Plots cleaned of pellets. Female calf introduced into 10-acre pen in Pen 2. Twenty traps complete: 11 inside pens; nine outside. Two male yearlings released from 10-acre pen into Pen 2. Replicate count experiments conducted. Pellet-plots counted and cleared in Pen 1. Browse utilization estimated in Pen 1. Numbers of moose in pens redetermined and calves counted, 10 Productivity and Mortality Mortality and natality within pens are measured by daily ground observations, periodic aerial observations, trapping and use of radio- tracking devices. General condition is estimated for trapped animals by methods described below. Blood Blood values are determined from serum and whole-blood samples obtained from trapped and hunter-killed moose and animals immobilized for marking outside of traps (Job 3). Table 2 lists sources of blood material. Blood is obtained from live immobilized animals in sterile evacuated containers by jugular venepuncture. Four or five cc of whole blood are preserved with EDTH and a thin smear is made; serum is secured by centrifugation of cooled and clotted blood, Serum is separated into: 1) a NaF tube (1.5-2 cc) for glucose determination; and 2) a 4-5 cc, untreated sample for other parameters. Analyses are performed by Alaska Medical Laboratories (Anchorage) using a Technicon Autoanalyzer SMA-12, standard hematological techniques and electrophoresis. Browse Production and Utilization Browse production and utilization and plant succession are estimated using methods•previously described in detail (Bishop, 1969). A canopy- cover method after Daubenmire (1959), employing exclosures, is used for successional measurements and a twig-count method with clipping is used for production and utilization estimates. During this reporting period use was estimated in all old plots in pen 1, the ten-acre holding pen. Weights and Measurements Weights and measurements were obtained from trapped immobilized animals. Tame Moose Feeding Feeding habits of tame moose were studied by standing about one meter from the moose and recording species and size of each bite eaten by pencil on an IBM optical page reader sheet or by speaking into a tape recorder. Bite size was recorded by number of leaves (estimated) in summer and by twig length (also estimated) in winter. Hours of observa- tion were distributed throughout daylight hours and periods of inactivity, urinations, defecations, breakage of shrubs and abnormal behavior were also recorded. Ninety-nine hours of observation of two males and one female (all aged 12-14 months) took place in a 10-acre holding pen during July and 11 Table 2. Sources of moose blood for analysis: June 1969-May 1971. NUMBER OF SPECIMENS Source Trapping at Moose Research Center: Pen 1 Pen 2 Pen 3 Pen 4 Outside pens Total Moose Research Center Hunts: 1969-70 GMU 15C 15B 14A 14B 1970-71 GMU 7 15A Total Hunts Tagging: Bottenintnin Lake (1970) Moose River Flats (1970) Skilak-Tustumena Beach (1971) Moose River Flats (1971) Total Tagging TOTALS 12 Serum 20 25 15 22 40 122 32 13 39 14 7 28 133 38 61 3 60 162 417 Whole Blood 15 16 13 15 26 85 26 6 26 6 2 0 66 23 0 0 0 23 174 Slides 5 10 7 12 9 43 49 21 42 9 0 0 121 6 0 0 0 6 170 August, 1970. Forty-one hours of observation of the same two males free- ranging in pen 2 and the same female confined to the 10-acre pen took place from February through May, 1971. Tame animals were obtained as calves in October 1969 (males) and May 1970 (female) from Dr. Jack Luick of the Reindeer Experimental Station at Cantwell. They had been raised on browse and artificial feed and were supplementally fed calf pellets until July 1970. The three were confined together until 11 November 1970, when the males were placed in pen 2. The female gave birth to a single calf on about 2 July 1971. Average bite sizes were converted to weight by collecting and weigh- ing 500 birch (Betula papyrifera) leaves (20 from each of 25 plants selected randomly) and twigs and whole plants representative (subjectively) of various bite sizes recorded during feeding observations. Food Habits from Rumen Analysis Rumen samples were collected during antlerless moose hunts in upland range (Juneau Creek; Unit 7; 2-6 December 1970; n = 8) and lowland seral range (Unit !SA; 15-16 December 1970; n = 8). One-liter samples of least- digested rumina were collected from these animals. The samples were taken from recently· killed animals (less than 24 hours) and placed in plastic bags. They were washed with water on a 10 mesh/inch soil sieve within the next 24 hours and the washed solid fractions were placed in jars with 5 cc's of 10 percent formalin, shaken vigorously and frozen. For subsequent analysis the procedure used by Martin and Korschgen (1963) and Brown (1961) was used. Each sample was thawed and washed over a 10 mesh/inch soil sieve. A subsample of approximately four tablespoonfuls (50-150 ml) of the material retained on the sieve was used for the detailed analysis. The unused portion was examined for species not found in the subsample and for parts of species that would aid in recognition of smaller parts in the subsample. The subsample was placed in a flat-bottomed pan and floated in about 1/2 inch of water. Individual particles were picked up with forceps, identified, and placed with others of the same species or typed into a small dish. Unknown species and plant parts were separated and ultimately identified by comparison to better preserved portions of the same species. Plants that could not be identified to species were identified to a main group such as gramineous plants, sedges, browse, etc. A reference collection of browse twigs and reference to Morris et al (1962) and Hulten (1968) also aided in identification of plant parts and species. When examination failed to show any more identifiable particles, the remainder of the sample was measured volumetrically by water displace- ment in a graduated cylinder, and listed as unidentifiable. Practically all unidentifiable material was believed to be browse. Over 90 percent (approx,) was woody material with all bark and other diagnostic parts removed. The remainder consisted of parts too small to recognize. 13 The separated particles were measured by the same volumetric method, and the volume of each species or plant type recorded. Quantities too small for measurement were recorded as traces. Rumen Protozoa Twenty cc. of rumen liquor were collected from 18 animals killed in the same hunts, strained through cheesecloth and the liquor was fixed with 0.25 cc. 10 percent formalin. A drop of liquor was diluted 1:10 with Lugol's iodine and shaken and then placed on a hemacytometer, where the protozoa in 10 ~ were converted to protozoa/ml. After thorough mixing, 5 cc. of rumen liquor was placed into each of three graduated centrifuge tubes. Tubes were spun for 15-20 minutes at maximum centrifuge RPM (Aloe International Clinical Centrifuge) and proportion of sediment was recorded by measuring the supernate in a syringe. Chemical Composition of Plants Birch, willow, lowbush cranberry and aspen (bark) were collected for nutritional analysis according to plant height, diameter and use- form class. All specimens were oven-dried at 70° C for 14 days. Snow Monitoring Seven snow plots were established in pens 1 and 2. One plot was placed in each of the following habitat types: dense hardwoods, thin hardwoods, sedge meadow, spruce regrowth, birch-spruce regrowth (thin), birch-spruce (dense), spruce-ledum. At approximate 10-day intervals, a trench was dug in each plot and thickness and general consistency (eg: crystals, powder, ice) of each snow layer was recorded. Visibility of lowbush cranberry (Vaccinium vitis ideae) was also recorded at this time. FINDINGS Productivity and Mortality Within Pens Table 3 presents raw tagging, breeding, and mortality data for all moose within the enclosures. Table 4 summarizes these data in terms of numbers calving and dying. Table 5 calculates natality, yearling recruit- ment and change in population size for penned and unpenned populations. Figure 2 compares trends of natality and recruitment inside and outside the pens, On 1 July, 1971, populations within the four pens totaled 75 moose (18.8 per square mile). Components of the populations are tabulated in Table 6. Although the matter is complicated by introductions, experimental collections and escapes, it appears that numbers have returned to their approximate level when the pens were first enclosed. Populations in each pen declined initially for various reasons (pen 1 because of escapes and 14 Table 3. Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex 1 F 2 F 3 F 4 F 5 F Event Tagged Pregnant With 2 calves With 1 calf Escaped into pen 2 with 1 calf Tagged With 1 calf Escaped into pen 2 with 1 calf Tagged Pregnant With 1 calf With no calf With no calf Radio installed Radio removed In oestrus With 1 calf (/1370UC) Weight, 920 pounds Calf survived to yearling With 1 calf Tagged With no calf, does not appear pregnant Dead, not pregnant Tagged Last seen (presumed dead) Date January 1968 January 1968 7 June 1968 17 June 1968 PEN 1 4 September 1968 January 1968 4 June 1968 January 1968 January 1968 17 June 1968 11,14 October 1968 2,11,12,14 June 1969 14 June 1969 20 August 1969 15 October 1969 2 July 1970 10 September 1970 3 June 1971 3 June 1971 January 1968 5 June 1968 18 May 1969 18 January 1968 18 January 1968 Age 4+ years 5 years 4+ years 5 years 5+ years 6 years 7 years 8 years 9 years 15+ years 16 years 17 years Circumstances Helicopter; not with calf Palpation Helicopter; not with calf Observed Helicopter; not with calf Palpation Observed Observed Observed and trapped Trapped Trapped Trapped Observed Trapped Observed (/1370UC) Observed Helicopter, not with calf Observed Apparent cause: old age Helicopter; not with calf Tagging ...... "' Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures~ Janu&cy 1968 through June 1971 Moose II Sex Event 6 F Tagged With calf With no calf Weight, 910 pounds With one calf (11670 Female) Calf survived to yearling No calf produced 8 F Tagged Dead 10 F Tagged With no calf With 1 calf {II1070UC) Weight, 725 pounds Calf survived to yearling With 1 calf R70-8 F Tagged, radio installed With 1 calf 40 F Five observations With 1 calf (4070) Tagged Calf survived to yearling With 1 calf PEN 1 Date January 1968 22 October 1968 2,4,11,12 June 1969 12 August 1970 23 July 1970 3 June 1971 3 June 1971 24 January 1968 2 February 1968 January 1968 19 June 1969 27 July 1970 6 August 1970 3 June 1971 3 June 1971 10 July 1970 3 June 1971 21 May 1969 through 30 June 1970 27,28 July 1970 1 September 1970 3 June 1971 3 June 1971 Age 10+ years 11+ years 12 years 13 years 14 years Calf Calf 2 years 3 years 4 years 2 years 3 years 2 years 3 years Circumstances Helicopter; not with calf Observed Observed, trapped Trapped Trapped, weight 820 pounds Observed Observed Helicopter Killed during tagging Helicopter Deserved Observed Trapped Observed Observed Trapped Observed Observed Observed Trapped Observed Observed Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex 41 F uc F 35 M 43 M 670 F 370UC M Event Broke into pen with 4170 (calf) Tagged Released outside pen as /183 Broke into pen With 1 calf Tagged Weight, 605 pounds Weight, 715 pounds Antler spread 77.5 em. Last seen Tagged & put into pen 1 Apparently did not successfully breed Bred successfully Right antler: 10 points 100 em. spread est. 120 em., weight 5 lbs. Bred successfully Last seen Born to /16 Female First observed Tagged Survived to yearling Born to /13 Female First observed Survived to yearling PEN 1 Date 13 October 1970 14 October 1970 23 February 1971 14 October 1970 3 June 1971 28 May 1970 17 July 1970 22 September 1970 7 March 1971 7 June 1968 May-June 1969 1969 1970 3 June 1971 May-June 1970 22 July 1970 23 July 1970 3 June 1971 After 21 May 1970 2 July 1970 3 June 1971 Age 7+ years 7+ years 2 years 2+ years 2+ years 1+ years 2+ years 3+ years 4 years Calf 1 year Calf 1 year Circumstances Observed, hole in fence Trapped Trapped Observed Observed Trapped Trapped Trapped Trapped Observed Helicopter immobilization No calves in pen 1, May-June 1969 Calves in pen 1, May-June 1970 Recovered Calves in pen 1, May-June 1971 Observed Observed Trapped Observed /13 Female observed pregnant Observed Observed I-' co Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex Event 1070UC M Born to 1110 Female First seen Trapped -escaped Survived to yearling 4070UC ? Born to 1/40 female First seen Survived to yearling Moose II Sex Event 1 F Escaped into pen 2 with 1 calf With no calf With no calf Weighed 675 pounds With no calf With no calf 2 F Escaped into pen 2 with 1 calf With 1 calf Calf not seen With no calf Radio installed Not pregnant ~-lith no calf With 1 calf PEN 1 Date May-June 1970 27 July 1970 15 September 1970 3 June 1971 May-June 1970 15 September 1970 3 June 1971 PEN 2 Date 4 September 1968 10 October 1968 15 July 1969 5 September 1969 10 June 1970 3 June 1971 4 September 1968 4 September 1968 21 April 1969 30 May, 7 July 1969 13 August 1969 28 May 1970 3 June 1971 Age Calf 1 year Calf 1 year Age 5+ years 6+ years 7 years 8 years 5+ years 5 years 6 years 7 years Circumstances Observed Trapped Observed Observed Observed Circumstances Observed Observed Trapped Trapped Observed Observed Observed Trapped Trapped Trapped Observed Observed Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II 7 (R70-7) 9 11 Sex F F F Event Tagged With no calf With 1 calf With 1 calf; weight 710 pounds In oestrus, with 1 calf With no calf Not pregnant, with no calf Radio installed Weight: 865 pounds With 1 weak, deformed calf Tagged With no calf With 2 calves With 1 calf With no calves Breeding (?) With no calves With 2 calves Tagged With no calf "Especially" poor condition With no calf In oestrus Last seen alive Carcass found PEN 2 Date January 1968 18 June 1968 9 July 1969 29 July 1969 23 September 1969 26,27 February 1970 4 June 1970 4 June 1970 23 July 1970 25 May 1971-7 June 1971 January 1968 5,17 June 1968 11 June 1969 12 June 1969 10 July 1969 31 October 1969 4,25 June 1970 14 June 1971 January 1968 21 May, 10 October 1968 1 May 1969 1 July 1969 24 September 1969 26 January 1970 3 June 1971 Age 4+ years 5 years 6 years 6+ years 7 years 7+ years 8 years 5+ years 6 years 7 years 7+ years 8 years 9 years Circumstances Helicopter; not with calf Observed Observed Trapped Trapped Helicopter; observed Trapped Trapped Trapped Observed Helicopter; with 1 calf Observed Observed; calves tagged Tagging-induced mortality Tagging-induced mortality With i/45 male Trapped, observed Observed Helicopter; not with calf Observed Observed Observed Trapped Observed Dead more than a year N 0 Table 3. {cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose IJ Sex 12 F Autopsy: 13 F 16 F 17 F 52 F Event Tagged Not with calf Not with calf Lost. Seen alive Found dead Massive hemmorage left of Lactating Flaccid uterus Weight: 610 pounds PEN 2 Date January 1969 21 May 1968 18 February 1969 1 May 1969 7 July 1969 rib cage; dead along Age 6+ years 7 years 7+ years 8 years 8+ years fenceline Circumstances Helicopter; not with calf Observed Observed Observed Died 3-6 July Conclusions: Death perhaps due to combination of poor condition and hitting fence. Tagged With no calf With 1 calf; wt. 580# Found dead Tagged With no calf With no calf Hock tumor & displasia Last seen (presumed dead) Tagged Found dead Tagged With 1 calf Calf survived to yearling With 1 calf January 1968 4 October 1968 30 July 1969 August 1969 January 1968 18 June 1968 9 July 1969 3 August 1969 26,27 January January 1968 25 April 1968 23 July 1969 18 June 1970 3 June 1971 7 June 1971 7+ years Helicopter with 1 calf 8+ years Observed Trapped 9+ years Drowned in hole 9+ years Helicopter with no calf 10 years Observed 11+ years Observed Observed 1970 11+ years Observed from helicopter Calf Helicopter Dart in carcass 2+ years Trapped 3 years Observed Observed 4 years Observed N 1-' Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center encLosures, January 1968 through June 1971 PEN 2 Moose II Sex Event Date Age Circumstances R70-2 F Tagged 22 May 1970 3 years Trapped Lactating; wt. 635 pounds 17 July 1970 Trapped Calf last seen (survival unknown) 4 September 1970 4 years R70-2 not seen until 3 June 1971 With no calf 3, 7 June 1971 Observed R70-4 F Tagged & introduced into pen 2 from outside 23 May 1970 3 years Trapped Deserts calf 23 May 1970 Trapping-induced desertion With 1 calf 3 June 1971 4 years Observed R70-5 F Tagged, introduced into pen 2, radio installed 24 May 1970 9 years Trapped Died giving birth 27 May 1970 Twin fetuses; one breech presentation Raquel F Brought from Cantwell as tame calf: In 10-acre pen May 1970 1 year Tame moose Confined with 2 1+ year old males Through 30 November 1970 1+ years With no calf 7 June 1971 2 years Observed Uncollared F Observed 1 June 1969 2 years(?) Observed Six observations Through 25 May 1970 Observed With 1 calf 8 June 1970 3 years(?) Observed from helicopter Last seen with calf (calf seen 21 Jan.-- survival probable) 30 September 1970 Observed With 2 calves 25 May-7 June 1971 4 years(?) Observed N N Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose ft Sex 3995 F 15 M 4250 M 36 M 45 M Walter M Event Tagged Dead Tagged Last seen alive Carcass found Tagged Last seen (presumed dead) Observed Breeding (?) Bred successfully Antler spread (velvet): green Breeding Bred successfully Last seen Tagged Antler spread (velvet): 23 em. Breeding condition Last seen Brought from Fairbanks as tame calf: In 10- acre pen Rutting Weight, 660 pounds Antler spread 71 em. PEN 2 Date January 1968 Smelled: 2 July 1969 Found: 8 July 1969 January 1968 10 October 1968 3 June 1970 January 1968 January 1968 23 July 1969 3,14,15 October 1969 Fall 1969 29 July 1970 Fall 1970 Fall 1970 3 June 1971 21 October 1969 5 April 1970 October 1970 3 June 1971 September 1969 October 1970 to November 1970 Age 5+ years 7 years 2+ years 3+ years Calf 2+ years 2+ years 2+ years 3+ years 3+ years 4 years 1+ year 2+ years 3 years Calf 1+ years Circumstances Helicopter; with no calf No autopsy possible Helicopter Observed Dead more than a year Helicopter Tagging Observed Observed with adult females At least 2 calves in pen 2 Trapped Observed with adult females in rutting behavior Calves in pen 2 May, June 1971 Observed Trapped-no antler development Trapped In rutting group--forced away by #36 male Observed Tame moose Observed Freeze-branded N w Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose /J Walter (cont'd.) Richard Sex M R70-2-70UC ? UC70UC ? 5270UC ? 5050 F Event Released into pen 2 with bell Last seen Brought from Fairbanks as tame calf: In 10- acre pen Weight: 690 pounds antler spread: 75 em. Released into pen 2 with radio Last seen Born to R70-2 First seen Last seen with cow Survived to yearling(?) Born to UC First seen Last seen with cow Survived to yearling(?) Born to 52 First seen Survived to yearling Tagged; with no calf Killed; in oestrus PEN 2 Date 30 November 1970 21 April 1971 September 1969 11 November 1970 30 November 1970 3 June 1970 22 May 1970 12 June 1970 14 September 1970 3 June 1971 May-June 1970 27 July 1970 30 September 1970 3 June 1971 May, June 1970 18 June 1970 3 June 1970 August 1969 23 October 1969 Age 2 years Calf 1+ years 2 years Calf 1 year Calf 1 year Calf 1 year 2+ years 2+ years Circumstances Observed Tame moose Freeze-branded Released Observed R70-2 trapped -placenta observed Observed Observed Two yearlings observed in pen 2 Observed Observed Two yearlings observed in pen 2 Observed Observed with 52 Trapped Overdose of MS0-50 diprenorphrine Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex 20 F 26 F Autopsy: 27 F 28 F Event Tagged; with 1 calf With 1 calf Calf survived to yearling (ti2069UC) With 1 calf Tagged; with 1 calf In oestrus With no calf Killed One fetus Total weight -600 pounds PEN 3 Date 6 August. 1969 27 February 1970 29 June 1970 3 June 1971 23 September 1969 29 October 1969 26,27 January 1970 19 May 1970 Weight less fetus & fetal membranes -560 pounds Tagged 26 September 1969 With 1 calf 8 October 1969, 27 January . With no calf; not lactating 10 July 1970 With no calf 3 June 1971 Tagged; lactating 6 October 1969 In oestrus With no calf 26,27 January 1970 With 1 calf; weight 760 pounds 17 July 1970 Weight 765 pounds 15 September 1970 Calf survived to yearling (112870) 3 June 1971 With no calf 3 June 1971 Age 9+ years 9+ years 10 years 11 years 8+ years 9 years 3+ years 1970 4+ years 5 years 7+ years 8+ years 9 years Circumstances Trapped Observed from helicopter Observed Observed Trapped Trapped Observed from helicopter Overdose M-99 Etorphine Trapped Trapped & observed from helicopter Trapped Observed Trapped Observed from helicopter Trapped Trapped Observed Observed Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 PEN 3 Moose If Sex Event Date Age Circumstances 38 F Tagged: not lactating 10 July 1970 16 years Trapped With no calf 3 June 1971 17 years Observed 39 F Tagged: with 1 large calf 28 July 1970 5+ years Trapped Calf survived to yearling ca: 20 May 1971 Observed With 1 calf 3 June 1971 6 years Observed uc F Observed 9,15 July 1970; 7 March 1971 1+ years(?) Observed N I.JI 60 M First seen 6, 7 Ocwber 1969 2+ years Observed, trapped Bred successfully Fall 1969 2+ years 2 calves in pen 3 May,June 1970 Tagged 15 May 1970 3 years Trapped Bred successfully Fall 1970 3+ years 2 calves in pen 3 May, June 1971 Last seen 3 June 1971 4 years Observed 2069UC M Born to 1120 May-June 1969 Calf First seen 12 August 1969 Observed Last seen with 1!20 14 October 1970 1+ years Observed Last seen 3 June 1971 2 years Observed 2870 F Born to 1!28 May-June 1970 Calf First seeu 17 July 1970 Observed Tagged; weight 246 pounds 12 August 1970 Trapped S u Y"Vi ve<1. to yearling 3 June 1971 1 year Observed J.J.JOUC C' Bc.rn to 1!39 May-June 1970 Calf First seen 17 August 1970 Observed Survived to yearling 3 June 1971 1 year Observed N "' Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures~ January 1968 through June 1971 PEN 4 Moose ff Sex Event Date Age Circumstances 22 F Tagged~ with calf August 1969 4+ years Trapped With 1 yearling 25 May, 8 June 1970 5 years Observed With no calf; not lactating; weight 660 pounds 17 July 1970 5+ years Trapped Weight 785 pounds 1 September 1970 Trapped With 1 calf 20,25 May; 3 June 1971 6 years Observed 23 F Tagged 4 September 1969 11+ years Trapped With 2 calves In oestrus With 2 calves 29 October 1969 Trapped Found dead 13 April 1970 11+ years Autopsy: No fat Flaccid vascularized empty uterus 24 F Tagged, no calf seen August 1969 7+ years Trapped With 1 calf 26 January 1970 Observed from helicopter With 1 calf 25 May, 2 July 1970 8 years Observed Calf survived to yearling 3 June 1971 Observed With 1 calf 3 June 1971 9 years Observed 25 F Tagged; with 1 calf weight 920 pounds 5 September 1969 10+ years Trapped With 1 calf 29 December 1969 Trapped Seen alive; with no calf 26, 27 February, 4 February Observed from ground & helicopter Found dead 20 July 1970 11+ years Dead before 1 June, cause unknown 29 F Tagged; with 1 calf 14 October 1969 5+ years Trapped In oestrus 14 October 1969 Trapped N -...J Table 3. (cant' d.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex Event 29 (cont 1 d) Last seen alive with 1 calf Counts Found dead 31 F Tagged with 1 calf (R70-l) With 1 calf Calf not seen Calf born Calf survived to yearling With 1 calf 34 F Tagged With no calf With no calf 36 F Tagged: with !'calf Calf survived to yearling With 1 calf 37 F Tagged: weight 585 pounds With no calf R70-3 F Tagged Cne calf born Weight: 825 pounds Calf survived to yearling With 1 calf PEN 4 Date Age 21 October 1969 26 January, 4 February 1970 23 April 1970 5+ years 12 August 1969 29 December 1969 26 January 1970 25-28 May 1970 3 June 1971 6 June 1971 11 December 1969 26 January 1970 3 June 1971 23 July 1970 ca: 25 May 1971 3 June 1971 1 October 1971 3 June 1971 20 May 1970 22-23 May 1970 1 September 1970 3 June 1971 3 June 1971 5+ years 6 years 7 years 13+ years 14 years 7+ years 8 years 1+ years 2 years 3 years 4 years Circumstances Trapped Not pregnant; dead more than 4-6 weeks in sitting position Trapped Observed Observed from helicopter Observed Observed Observed Trapped Observed Observed Trapped Observed Observed Trapped Observed Trapped Observed Trapped Observed Observed N co Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose I! A60 21 44 Sex Event F Tagged; no calf seen Not lactating Weight 840 pounds With 1 calf M Tagged Bred successfully Last seen M Tagged; large staph infection lwdt buttock PEN 4 Date 17 July 1970 3 June 1971 August 1969 Fall 1970 3 June 1971 9 October 1969 Not seen during aerial counts Age 13+ years 14 years 1+ years 2+ years 3 years 1+ years or after (presumed dead) 26 January-4 February 1970 7 M UC M 24 70UC ? 3670UC ? R70-l-70UC ? Tagged Last seen Observed Observed Born to 1124 First seen Survived to yearling Born to /136 First seen Survived to yearling Born to R70-l Survived to yearling 4 June 1970 3 June 1971 4 September 1970 3 June 1971 ca: May 1970 25 May 3 June 1971 May-June 1970 27 July 1970 3 June 1971 3 June 1970 3 June 1971 1 year 2 years 1+ years 2 years Calf 1 year Calf 1 year Calf 1 year Circumstances Trapped Observed Trapped Calves in pen 4; May-June 1971 Observed Trapped Trapped Observed Observed Observed Observed Observed Observed Observed Observed Observed Table 3. (cont'd.) Histories of individual moose in Kenai Moose Research Center enclosures, January 1968 through June 1971 Moose II Sex R70-3-70UC ? NB 4170 M Event Born to R70-3 Survived to yearling PEN 4 Date 23 May 1970 3 June 1971 Age Calf 1 year Circumstances Observed Observed Three dead calves/short yearlings were discovered in pen 4: one died on 14 February 1970 and was discovered that day. One apparently died during winter before 22 April 1970 near the carcass of #39 female. The third died probably during early April 1970. Six calves were present in the pen the summer and fall of 1969. Four of them were offspring of females that died during the 1969-1970 winter (#23, #29, and #25). Three yearlings (#7, #U/C, #37) survived the winter. Therefore, at least one yearling must have survived a winter during which its mother died. Born to #41 outside of pens Broke into pen 1 with 1141 Tagged, placed in pen 4 Probably did not survive to yearling May-June 1970 13 October 1970 23 February 1971 June 1971 Calf /141 Trapped Trapped Not observed since 23 February 1970 w 0 Table 4. Moose natality, mortality and recruitment in four one-square mile enclosures. Adult FF January 1968* 6 June 1968 6 June 1969 4*** June 1968-June 1969 June 1970 5 June 1969-June 1970 June 1971 6 June 1970-June 1971 January 1968* 8 June 1968 8 June 1969 11*** June 1968-June 1969 June 1970 10**** June 1969-June 1970 June 1971 9 June 1970-June 1971 * Ignoring tagging mortality id; Introduced into pen {MM) (0) (1)** (1) (2) (2) (1) (1) (1) (2) (4) Calves Calves Lost PEN 1 1 5 0 1*** 4 0 5 4 4 PEN 2 3 1 4 0 3 4 8 0 *** 2 Calves + 2 cows escaped into pen 2 Sept. 1968 **** One adult female introduced May 1970 Yearlings Adults Net Gain (+) or Loss (-) Recruited Died of Adults (Including (discounting experimental long yearlings) manipulation) 1 2 2 1 +1 0 0 0 0 +4 (1 female broke 0 00 000 4*** 1 1 3 +1 0 0 2000 2-3 2-3 2 2 Adult females killed One killed female contained 1 fetus One adult trapped in man-made hole -1 0 in) (2 males, 1 female introduced) w I-' Table 4. (cant 'd.) Moose natality, mortality and recruitment in four one-square mile enclosures. Adult FF August 1969 8 June 1970 60 August 1969-June 1970 June 1971 6 June 1970-June 1971 August 1969 12 June 1970 9 August 1969-June 1970 June 1971 8 June 1970-June 1971 * Ignoring tagging mortality ** Introduced into pen (MM) (1) (1) (2) (2) (1) (3) Calves Calves Lost PEN 3 4 zoo 1 2 PEN 4 10 4+ 5-6 6 0 *** 2 Calves + 2 cows escaped into pen 2 Sept. 1968 **** One adult female introduced May 1970 Yearlings Adults Net Gain (+) or Loss (-) Recruited Died of Adults (Including (discounting experimental long yearlings) manipulation) 0 3 2 (killed) 3 +1 2 2 2 0 1 4 4 4 -3 4 4 3 +1 0 2 Adult females killed 00 000 One killed female contained 1 fetus One adult trapped in man-made hole Table 5. June calf crops and yearling recruitment in Moose Research Center enclosures. Calf Crop Yearling Recruitment Population (Adult) Year Calves/100 F (No. F) Yrlgs/100 F (No. F) % Gain/loss (excluding (No. 1+ years manipulations) old) Pen 1 1968 83 (6) 17 (6) 1969 0 (4)(no breeding bull) so (4) +17% (6) 1970 100 (4) No recruitment No change (7) 1971 83 (6) 80 +52% (12) x (not incl. 69-70) 88 47 +38% w Pen 2 N 1968 12.5 (8) 25-38 (8) 1969 50 (8) 9 (11) +11% (11) 1970 30 (10) 0 (11) -8% (12) 1971 100 (8) 25-38 (8) No change (13) X 41 13-18 Pen 3 1969 (August) 50 (8) 0 (8) No data 1970 29 (7) 38 (8) No change (10) 1971 33 (6) 33 (6) No change (10) X 38 23 Table 5. (cont'd.) June calf crops and yearling recruitment in Moose Research Center enclosures. Year Pen 4 1969 (August) 1970 1971 X All Pens 1968 1969 1970 1971 X Calf Crop Calves/100 F 83 45+ 75 69+ 43 59 43 72 56 (No. F) (12) (9) (8) (14) (27) (30) (29) (100) Yearling Recruitment Yrlgs/100 F (No. F) 8 44-56 50 31-34 21-29 11 25-29 41-45 2 25-27 (12) (9) (8) (14) (35) (28) (29) (106) Unit 15A (Aerial counts by Richey (unpublished) and LeRoux (unpublished) and LeResche) 1968 1969 1970 1971 X 47 48 ca: 30 21 39 (1520) (438) (ca: 500) (657) (ca: 3115) 18 (est.) 7 (est.) 23 14 17 (1520) (438) (496) (166) (2620) Population (Adult) % Gain/loss (excluding (No. 1+ years manipulations) old) No data -21% + 7% -7% +13% -10% +11% + 3% + 4%** (14) (15) (15) (39) (44) (98) ** Kenai National Moose Range stratified random mile-square quadrant counts (unpublished) indicated populations north of the Kasilof River of 6700 ± 1410 in winter 1967 and 7900 ± 1460 in winter 1971. This represents a mean annual increase of 04+ % for the four years, disregarding the variations in weather and observers and ignoring the confidence intervals. Figure 2. June calf and yearling crops inside and outside Moose Research Center enclosures, 1968 -1971. I~ Within Pens :z ::l Q--Outs i de Pens -, • Of-. g 34 ' ' ' ' ' ----o f(j7 Table 6, Populations within Kenai Moose Research Center enclosures as of 14 June 1971. Females with No Calves 1 Calf 2 Calves Yearlings Males Total Pen 1 1 5 0 4 2 17 Pen 2 2 5 2 (2):3 4 25 Pen 3 4 2 0 2 2 12 Pen 4 2 6 0 4 3 21 All Pens 9 18 2 13 11 Moose 9 36 6 13 11 75 Summary FF Yrlgs Calves MM Total Pen 1 6 4 5 2 17 Pen 2 9 3 9 4 25 Pen 3 6 2 2 2 12 Pen 4 8 4 6 3 21 All Pens 29 13 22 11 75 35 lack of available bulls; pen 2 by collection and mortality; pen 3 by human-caused mortality; pen 4 by mortality), to the levels of February 1970 (see Methods Section). Upon closure, the pens contained 16, 24, 12 and 18 moose (total= 70), respectively, by best counts. This agrees approximately with a population of 75 immediately following calving. Probability of each animal's dying seems to peak in the first year after its enclosure in a pen (Table 3). Animals remaining after that time seem to have adapted to the relative confinement and to more closely follow norms for unenclosed moose. Of 12 adults (greater than 24 months of age) dying naturally, eight died after being in the pens less than one year. As reported (LeResche, 1970), calf production and yearling recruit- ment for the years 1968-1970 were similar inside and outside the pens (Tables 4, 5; Fig. 2). During the past year (July 1970-June 1971) the relationship appears to have altered. Calf production and yearling survival were both much greater within the pens than outside (73 vs. ca:25 calves:lOO cows and 45 vs. 14 yearlings:lOO cows). Whether the differences are artifacts of small numbers (only 29 adult females are present in the pens), are due to imprecise sampling outside the pens, or represent a 'true biological response remains to be seen. The possibility that increased calf production and yearling survival resulted from the temporarily depressed population inside the pens is an intriguing one. Overall adult population change within the four pens since 1968, eliminating experimental alterations, was +3 percent (Table 5). Annual changes ranged from +13 percent (1968-69) to -10 percent (1969-70). Outside the pens, the only estimates of moose numbers are from stratified random mile-square quadrant counts accomplished by Kenai National Moose Range personnel in winter 1967 and again in winter 1971 (unpublished). In 1967, the population north of the Kasilof River was estimated at 6700 ± 1410 moose. In 1971, the count estimated 7900 ± 1460 in the same area. Disregarding variations in weather, counters, etc., and the confidence intervals, this represents approximately a four percent annual increase for the four-year period. Thus, there is the secondary possibility that increases within the pens are truly reflecting outside changes~ LeResche (1970) tabulated mortalities within the pens through July 1970. Table 7 updates this information to include mortalities and carcasses discovered since then. Both calves and animals 12-24 months of age died in numbers greater than expected from their proportions in the population. Seven of 25 nonhuman-related mortalities (28 percent) were calves, whereas during the three years calves comprised only 23/107 (21 percent) of the population (difference not significant; P) .20; t-test). Similarly, animals 12-24 months old represented 6/25 (24 percent) of recorded mortalities, while making up only 15/107 (14 percent) of the population (P ( .10; t-test). In contrast, adults comprised 48 percent of the mortalities while making up 55 percent of the population during the period. During 1970-71, no adult mortalities occurred. During the same period six moose 12-24 months old died, as did one calf, which had been 36 Table 7. Mortalities within pens January 1968 -June 1971. MOose # Sex Age Pen # 4 F 17+ l 5 F Ad 1 11 F Ad 2 12 F 8+ 2 13 F Ad 2 14 F 9+ 2 15 M 3+ 2 Month -Year April 1969 January 1968 After January 1970 May-July 1969 After June 1968 August 1969 After October 1968 16 R70-5 F F 11+ 2 After January 1970 3995 5050 26 P3691 P3692 23 25 29 44 P4691 P4692 P4693 8 17 4250 4170 368 168 769 969 969 1269 25(69) 2969 GM69 F F F ? ? F F F M ? ? ? F F M M ? ? ? M F ? M ? ? 9 Outside: put in pen 2 May 1970 7 1+ 9 1+ 1+ 11+ 11+ 5+ 1+ 1+ 1+ 1+ Calf Calf Calf Calf Calf Calf Calf Calf Calf Calf Calf Calf Calf 2 3 3 3 3 4 4 4 4 4 4 4 1 '2 2 1-4 1 1-2 2 2 2 2 4 4 4 July 1969 October 1969 May 1970 Nov. 1970-June 1971 Nov. 1970-June 1971 Oct. 1969-Apr. 1970 Feb. 1969-May 1970 Oct. 1969-Apr. 1970 Oct.-Dec. 1969 Feb. 1970-Mar. 1971 Feb. 1970-Mar. 1971 July 1970-Mar. 1971 January 1968 January 1968 After Jan. 1968 After 23 Feb. 1971 June-October 1968 Sept. -Oct. 1968 January 1970 June 1969 June 1969 July 1969 1 February 1970 Oct. 1969-Jan. 1970 March-April 1970 37 Cause Winter -old age: carcass found Never seen after tagging Winter-kill. Carcass found Unknown. Carcass found Unknown. Disappeared Drowned in hole. Very poor condition Unknown. Carcass found with antlers intact Unknown. Disappeared Calving complications, carcass found Unknown, carcass found Killed with drug Killed with drug; pregnant; wt. 600 pounds Unknown, disappeared Unknown, disappeared Unknown; had aborted; no fat; carcass found Unknown; carcass found Unknown; carcass found; not pregnant Disappeared; badly infected rump when last handled Unknown; disappeared Unknown; disappeared Unknown; disappeared Killed during tagging Killed during tagging Never seen after tagging Never seen after put into pen 4 Unknown; disappeared Unknown; disappeared Unknown; carcass found Deserted after tagging Deserted after tagging Mother died July Pneumonia; wt. 160 pounds Mother died (see above) Unknown; carcass found experimentally removed from its mother on 23 February. Although mortality in the 12-to 24-month-old age group (rather than before 12 months) is surprising, shift of mortality to younger age groups does indicate accli- mation of adults to the pens, as well as implying that maximum sustainable numbers of adults are present. Age structures of the penned populations are statistically equivalent (x2 == 36.7; df ==51; P) .10) to one another. Ages of male penned animals retain characteristics of hunted surrounding outside populations, for no penned bull is greater than four years of age. Age structures of all penned populations are statistically equivalent (x2 = 26. 4; df = 17; P ) .10) to those of animals trapped outside the pens. Blood Values as Indicators of Nutritional Status Specimens of blood analyzed through May 1971 are listed in Table 2. In addition, sera from 12 winter-killed animals from Fairbanks (collected by J. Coady) were analyzed. Seasonal, sexual and age-specific and pregnancy means of about 250 samples were reported previously (LeResche, 1970). The current data are presently being analyzed using factor-anlaysis and other multi-variant methods. The analytical scheme relates each parameter to several others and to age, sex, hind foot, total length, girth, girth/total length ratio, weight, date (season), reproductive status, location, and method of collection of the blood. Preliminary electrophoretic analyses were not reported in 1970. Table 8 lists seasonal means for albumin, total protein and albumin/globulin ratio for samples collected since then. Results suggest a decrease in TP, albumin and H/G ratio during the winter months. A more thorough analysis is being compiled using the above analytical program. Fig. 3 compares eight blood values of the 12 Fairbanks winter-killed specimens with early-winter means from Kenai (LeResche, 1970 and this paper) assumed to represent "normals." Though certainly inconclusive, Fig. 3 suggests results to be expected from the detailed analyses described above. Winter-killed animals had the following blood values: Calcium 9.1 ± 2.0 SD mg% (90% of "normal"). Phosphorus 5.7 ± 2.3 SD mg% (83% of "normal"). Calcium and phosphorus levels decrease with decreased intake (asso- ciated with hypoproteinemia) and vitamin D deficiency (LeResche, 1970). Glucose 54± 42 SD mg% (52% of "normal"). Glucose depletion usually accompanies terminal starvation, but values are maintained at near normal even in severe starvation. Possibly, "normal" mean from trapped and drugged animals is actually abnormally high and this lower value in winter-kills is closer to that found in undisturbed animals. In bovines and ovines, values vary from 35-74 mg% (Coles, 1964). 38 Table 8. Seasonal means (mg%} for total protein, albumin and albumin- globulin ratios in moose sera. Kenai Moose Research Center. Season n Total Protein Albumin A/G Ratio Rut 27 7.99 ± 0.69SD 5.07 ± 0.50SD 1.77 ± 0.304SD Early Winter 43 6.49 ± 0.96SD 3.76 ± 0.55SD 1.45 ± 0. 28SD Spring 9 7.54 ± 0, 70SD 4.66 ± 0.51SD 1.63 ± 0.28SD Summer 23 7.90 ± 0, 89SD 4.98 ± 0.69SD 1. 79 ± 0.45SD 39 - ,_ Figure 3: Selected blood values of winter-killed moose in relation to norms for the same time of year. t-.,00 2Cn r-100 ~ ~ ~I' ~lq ~ ,-I S> 40 Undisturbed winter-killed moose best approximate this range. Blood urea nitrogen 20.5 ± 12.6 SD mg% (340 percent of "normal). Preliminary results (LeResche, 1970) were ambivalent, in that highest values were recorded in spring but mean values decreased progressively from summer through late winter. The former suggested elevations in low nutrition, possibly due to protein catabolism; whereas, the latter pattern suggested values falling concurrently with nitrogen intake. Houston (1969) observed something similar to the latter in 12 moose. Current results (elevation of BUN in starvation) suggest that BUN produced by protein catabolism caused an elevation greater than the depression caused by decreased nitrogen intake. Dror et al (1970) and others have reported a negative correlation between nitrogen retention and BUN values, indicat- ing that poorly digestible foods were correlated with elevated BUN values. Oh ~ al (1969), in contrast, showed that BUN levels were elevated in sheep on low quality range forage when the animals were fed supplemental urea, and that BUN levels were significantly and positively correlated with rumen gas and VFA production and with dry matter intake. The apparent elevation of BUN in starving moose is only partially understood at present, therefore. Uric Acid 0.25 ± 0.15 SD mg% (10% of "normal). Theoretically, uric acid should be elevated in starvation (LeResche, 1970). The apparent decrease in winter-killed animals is unexplained at present. Total protein 5. 8 ± 1. 3 SD mg% (85% of "normal"). Albumin 3. 3 ± 0. 8 SD mg% (76% of "normal). A/G ratio 1.31 ± 0. 29 SD (87% of "normal"). Total protein, albumin and A/G ratio could all be expected to decrease in starvation, with albumin being the most sensitive indicator (LeResche, 1970). Present results follow the expected pattern. Browse Production and Utilization Studies of browse utilization using previously described methods (Bishop, 1969) proved to be too imprecise for practical use, even with relatively large sample sizes (LeResche, 1970). This portion of the job was therefore subordinated during this period as U.S. Fish and Wildlife Service personnel began to redesign the sampling procedure. R. Seemel and J. Oldemeyer did estimate utilization in pen 1. Results are given in Table 9 for comparison with previous years' data for the same plots (Bishop, 1969; LeResche, 1970). Utilization was also estimated on 38 plots in the tame moose holding pen (Table 10). This approximately ten-acre pen (including one ca:four- acre lake) was protected from moose browsing from August 1967 until October 1969, when two calf moose were introduced. These two occupied the pen during the winter of 1969-70, with supplemental feeding. A third short 41 Table 9. Numbers of birch sterns browsed by habit at type. Pen 1, June 1971. Kenai Moose Research Center. · Birch Regrowth Mature Hardwoods Dense Medium Thin Spruce-Birch Spruce Dense Thin n 25 24 26 23 19 20 22 x stems browsed 69.7 61.3 38.4 24.1 1.1 0.8 11.4 S.D. 53.2 42.6 35.5 29.6 3.69 2.68 20.8 Table 10. Numbers of birch stems browsed and broken in a 10-acre holding pen. June 1971. Kenai Moose Research Center • .p. N Birch Regrowth Dense Medium Thin Spruce-Birch Spruce Stems browsed: Number of plots 3 5 13 9 8 x stems browsed per plot 53.5 118.0 37.0 43.1 59.8 S.D. 95.5 116 23.5 32.1 64.5 Stems broken: Number of plots 3 5 13 9 8 x stems broken per plot 4.5 1.4 2.3 1.55 1.62 S.D. 5.75 1.95 2.6 2.2 2.8 yearling was introduced in May 1970 and the first two were removed in early November 1970. Only one moose (18-24 months old) occupied the pen during the winter of 1970-71. This moose ate much Vaccinium and lichens, and number of browsed twigs in birch habitat types from the winter of 1970-71 is no greater than in pen 1. However, browsing was severe by comparison in spruce and spruce-birch types. This suggests that birch utilization in birch types might well be maximal in most of the birch habitat of the 1947 burn, since overstocking did not increase measurable use. Although broken stems per unit area (Table 10) were not estimated for habitat outside the holding pen, the number within the small pen (2.2 per plot or 490 per acre) is much greater than that in habitat supporting lower moose densities. Broken stems per acre may be an indicator of moose utilization fot this habitat, therefore. Results (Table 10) show that this small pen received the heaviest use yet recorded and other data (tame moose feeding studies, this report) indicate the range's depleted state by May 1971. Weights and Measurements Table 11 lists forty-one whole body weights of moose trapped and immobilized. Fig. 4 illustrates the relationships between weight and age and time of year. Fig. 4 suggests that growth in weight possibly continues throughout a moose's lifetime, with major seasonal fluctuations. Mean September (maximum for the year) weights increased from 700 pounds (318 kilograms) for 2+ year-olds to 840 pounds (382 kilograms) for 5+ year-olds and 900 pounds (409 kilograms) for animals over 10 years of age, Basal (June) weights increased similarly, but were 15-30 percent (100-250 pounds) lower than maximum weights. Sequential weighings of individuals, though complicated by changes in reproductive status (Table 12) and possibly in rumen weights (Ledger, 1968)', confirm seasonal changes. These sequential weights are connected by dashed lines in Fig. 4. Verme (1970) also documented seasonal weight fluctuations (of 15-24%) in moose, but believed they reached maximum weight by 2-1/2 (females) or 3-1/2 (males) years of age. In contrast, Jordan~ al (1971), after reviewing old literature, assumed full body growth was not reached until 6 (females) and 11 (males) years of age. They reported (again, from literature) seasonal weight fluctuations of less than seven percent for full-grown females. Our data suggest longer growth periods than either of these authors, and also indicate greater seasonal fluctuations in weight. Both these factors suggest crowded conditions (cf: Klein, 1970) and agree with our estimates of moose densities in excess of 15 per square mile in the 1947 Kenai burn (cf: above). Unfortunately, there are no other weight data yet compiled for Alaska moose; therefore, it is impossible to compare 43 Table 11. Forty-one whole weights of moose of known age, sex and reproductive status. Kenai Moose Research Center. 1969-1971. Date 12 August 29 December 1 February 23 February 25 February 8 July 22 September 1 October 10 November 11 November 17 July 22 September 23 February 25 February 19 May 17 July 17 July 1 September 29 July 23 July 7 July 30 July 28 July 17 July 15 September 17 July 6 August 1 September 12 August 23 July 22 September 2 September 5 September 9 September 10 September 1 September 22 September 21 October 15 October 22 October 14 October Moose 2870 8170 P469 4170 8570 SL 76 37 WA RI 35* 35* 82 85 26 R70-2 22* 22* R70-7* R70-7* 12 14 DM 28 28 A60 10 R70-3 6* 6* 6* R70-4 1 72 3 20 75* 75* PASS 79 56 Pen 3 0 4 1 0 0 0 4 2 2 1 1 0 0 3 2 4 4 2 2 2 2 0 3 3 4 1 4 1 1 1 2 2 0 1 3 0 0 0 0 0 Sex F M M M F M F F M M M M F F F F F F F F F F F F F F F F F F F F F F F F F F F F M Age 2-5 months 7 months 8 months 9 months 9 months 13 months 16 months 16 months 17 months 17 months 25 months) 26 months) 5+ years 8+ years 9 years 3+ years 5+ years) 5+ years) 6+ years) 7+ years) 8+ years 8+ years 8+ years 8+ years) 8+ years) 13+ year;'J 3+ years 3+ years 12+ years) 13+ years) 13+ years) 3+ years 6+ years 8+ years 8+ years 10+ years Adult Adult Adult 13+ years 4+ years Reproductive Condition and Remarks Weight (lbs.) Survived to yearling Died of pneumonia Died within 3 months "Tame" -hand reared ,;Tame" -hand reared 110 lb. gain in 36 days (3.1 lbs/day) With no calf With one calf One fetus; moose killed with normal drug dose With 1 calf, calf probably did not survive to yearling. Did not calve following spring. With no calf; gain, 125 lbs/45 days Conceived; (2.8 lbs/day) 1 Calf; did not survive to yrlg. No calf produced Dead; had lost calf recently Dead; with 1 calf Calf recently lost (?) With 1 calf-survived to yearling With 1 calf-no calf next spring With no calf-did not produce calf next spring With 1 calf-survived to yrlg-10 conceived With 1 calf-survived to yrlg-70-3 conceived No calf-produced calf next spring One calf-survived to yrlg Did not produce new calf With no calf(lost in May)-Produced calf next spring With no calf-did not calve next spring With 1 calf With 1 calf-calf survived-3 produced calf next spring With no calf-produced calf next spring With no calf-90 lb loss/29 days With no calf (4.3 lbs/day) Unknown With one calf Rutting 246 385 160 340 350 550 580 585 660 690 715 850 820 600 635 660 785 710 865 610 580 880 760 765 840 725 910 820 870 710 675 850 920 875 960 870 910 970 1050 * Indicates serial weights of an individual. 44 Figure 4. Whole body weight by age of Kenai Peninsula moose. Kenai Moose Research Center. 1969-1971. (Dotted lines connect sequential weights of individuals.) 0 0 e q ___ Female with calf Female without calf Male ---------Sequential weights of the s arne ani mal dJ l-._.... ___ _ t;~ I ~' J ---+-- 1- :I <!1 6 e Gi--..,...._-+----3 e i .. 4- 12 e I I I I • .a4 0 :!JE, 9 I I I I 0 0 I 0 0 p I I I 0 0 0 0 0 0 IDa Table 12. Weight data suggesting the nutritional "cost" of calving and rearing young. Weight Moose Age Date Circumstances (lbs.) 6 '12+ 12 August With no calf 910 6 13+ 23 July With one calf 820 6 13+ 22 September With one calf 870 (Calf rearing "cost" ca: 75/910 pounds. or 8 percent of total body weight in mid -August) R70-7 R70-7 6+ 7+ 29 July 23 July With one calf With no calf 710 865 (Calf rearing "cost" 155/865 pounds or 18 percent of total body weight in late July) 22 5+ 17 July With no calf ) 19 % 660 22 5+ 1 September With no calf )+ " 785 28 8+ 17 July With one calf ~+0% 760 28 8+ 15 September With one calf 765 6 13+ 23 July With one calf ~+6% 820 6 13+ 22 September With one calf 870 (22 gained 125/660 pounds = 19% of mid-July weight by September 28 gained 5/760 pounds = 0% of mid-July weight by September 6 gained 50/820 pounds = 6% of July weight by September By not rearing a calf, 22 "gained" 16% of mid-July weight, or 106 pounds.) 46 maximum weights of this dense population with others. Verme's (1970) Alces americana females weighed a maximum of 850 pounds; whereas, our largest cow weighed 970 pounds (445 kilograms) and nine of our 12 females over four years of age weighed more than 850 pounds in August-october. Table 12 lists sequential weights of cows with and without calves, and calculates the implied cost of calf-rearing in terms of weight loss. Of course other variables than a calf's presence or absence were probably involved, but the three cases outlined suggest that calf rearing "costs" 8-18 percent of a cow's July-August weight. Skuncke (1949) noted similar differences in weight between cows with calves and cows without calves. Morphometric measurements (total length, hind foot, girth, height at shoulder, ear and tail) were taken of most moose handled. Such measurements are assumed to be indicators not only of growth but also of condition (Klein, 1968). In order to select the measurements not indica- tive of condition, sequential measurements of individuals were tabulated and examined. Ear and tail measurements were discarded because they varied little if at all between individuals. Height at shoulder was discarded because it is difficult to measure accurately in a eternally recumbent animal and hence varied in a patternless way in sequential measurements of individuals. Hind foot was found to vary a mean of 4.2 percent in sequential measurements; total length 2.2 percent; and girth 16.4 percent (in a seasonally-consistent manner). Therefore the ratio of the most seasonally variable (girth) and most stable (total length) measurements was taken as the most probable nutritional indicator. Fig. 5 plots girth/length (G/L) ratios and weights for all animals weighed. Correlation coefficient for the relationship is +0.34 (P) .10), indicating that G/L ratio is not a valid indicator of weight for animals of all ages and weights lumped. It is, therefore, probably not a valid condition indicator, either, for many animals lumped. The four dashed lines in Fig. 5 connect weights and G/L ratios for individuals weighed and measured sequentially. In three of the four cases, G/L ratio was positively correlated with weight. In the fourth instance, total length measurement changed from 303 centimeters to 281 centimeters (7.3%) from the first handling to the second a month later, indicating an inaccurate measurement. Thus, for individual animals, changes in G/L ratio may be a valid indicator of change in weight. Fig. 6 plots seasonal changes in G/L ratios for seasonally measured (but not weighed) individuals. The figure indicates that G/L ratios fluctuate seasonally just as do weights (Fig. 4). The ratios reach a minimum in May-June and maximize in September-october, as does weight. Tame Moose Feeding Habits Close observations of tame moose revealed winter and summer differ- ences in time budget and hourly forage consumption, but showed little differences in quantity of food consumed during each hour of active feeding. 47 0·?0 0 -~ ~ ::c 0-bO co ~ 2 UJ -J ~ tl -~ Figure 5. Relationship of girth:length ratios to whole body weights of 20 moose. Kenai Moose Research Center, 1969-71. Dotted lines connect values for the same individuals. 0 ~ E)_, E) 0 \ \ ~ 0 \ \ E)\ .,.,../ 0 e ~ \ ,..,,..,...,. 0 ~,.,... e ,.,... .,.,. .,;"' 0 ,;"' ,/" I ~ 0 ~ G) WEltAMT IN POUNDS. SIOO Ci> 0 0 FE"MALES -MALES I~ ~ ~ ~ t&J _, J: tz -\!) Figure 6. Seasonal variation of girth:length ratio in seven individual moose. Kenai Moose Research Center, 1970-71. 0 FEMALES e ~L-t.S 70 ~' "', ~ "" ......... ' " ' " 60 " ' ' JAN MAR 49 JUL 0 I I I I d SEP NOV Fig, 7 and Table 13 show concentration of feeding periods into reduced daylight hours during winter. During summer (July-August) feed- ing activity peaks occurred between 0400 and 0600 hours, when animals fed 89 percent of the time they were observed, and between 1600 and 2000 hours, when feeding occupied about 55 percent of the time. Feeding occurred for less than 43 percent of the time during all other daylight hours. In contrast, only one activity peak was evident in winter (February-May), but feeding occupied more than half the time during all daylight hours observed (Fig. 7). Forage consumption (bites per hour of day) was similarly concentrated in winter (Fig. 8; Table 14). Peak consumption in summer was 508 bites per hour between 0400-0800 hours. More than 350 bites per hour were con- sumed only during this period and between 1600-1800 hours in summer, when 380 bites per hour were eaten. During summer, moose averaged 237 bites per daylight hour, or 4270 bites per day (on a basis of 18 hours of daylight). By contrast, in each daylight hour during winter, moose consumed more than 350 bites, averaging 381 bites per hour and 4572 bites per 12-hour daylight period. Thus, number of bites consumed per day increased by seven percent in winter (4572-4270) but all consumption occurred in 4270 only two-thirds the time during winter. A good part of the increased bites per daylight hour in winter is explained by the greater proportion of time spent feeding during winter (Fig. 7; Table 13). This proportion increased from a mean of .38 to a mean of .64, a 68 percent increase during winter. In addition, bites consumed per hour £f feeding increased slightly in winter (Fig. 9; Table 15). In summer, moose averaged 646 bites per hour of feeding. In winter, consumption increased by 13 percent, to 730 bites per hour of feeding. Thus, increase in forage consumption (bites) per daylight hour and per day in winter may be attributed 84 percent ( 68 ) to increased 68 + 13 activity during each daylight hour and 16 percent to increased intensity of feeding during each active hour. The increase in mean bites per day by seven percent (from 4270 to 4572) from summer to winter is probably insignificant because of a change in data collection procedure and observer. Summer data were all recorded on IBM optical page reader sheets with a pencil. This required glancing from moose to sheet and may have caused missing some bites. Most winter data were spoken into a magnetic tape recorder and few bites, if any, were missed. Food habits varied significantly between summer and winter and moose ate a greater variety of forage during all seasons than was previously thought. Previous studies of deer species (Lay, 1964; Cowan~ al, 1970; Cushwa et al, 1970) have stressed that variety is important in diet and 50 Figure 7. Proportion of time spent feeding during each of 9 daily two-hour periods. 148 hours of observation of three moose. July and August, 1970. Kenai Moose Research Center. -~ 1·00 1\J: --·75 -~ • 5::::) -1-·25" -~ -~·75 -~·6'6 -~·25 11 J.IRS. 12 12 6 14 I~ I+ 8 4- JULY -AUGUST I I N'; 2.0 13 q.s 5·4 I FE'BR!lARY-fV\A. V I I I I 4 ' 8 10 l2 14 lb 18 2D 22 ~ ------~OU~ 0~ DAY----- 51 V1 N Table 13. Proportions of two-hour tline periods spent feeding by two male and one female tame moose, 99 hours of observation, July and August 1970 and 41 hours, February-May 1971. Kenai Moose Research Center. Time Period (ADT-hrs.) July-August 4-6 6-8 8-10 10-12 12-14 14-16 16-18 18-20 20-22 Hours feeding 12.5 5.0 1.0 0.5 5.5 5.5 7.5 4.5 0.25 Hours observed 14 12 12 6 14 15 14 8 4 Proportion of time spent feeding .89 .42 .08 .08 . 39 • 37 .54 .56 .06 February-May (AST-hrs.) 8-12 12-14 14-16 16-18 18-20 Hours feeding 10.8 9.2 5.9 4.7 0.6 Hours observed 20.2 13.3 9.5 5.4 1.0 Proportion of time spent feeding .53 .69 .62 .87 .60 Figure 8. Total consumption of natural forage by three tame yearling moose in relation to time of day. 99 hours of observation in July and August, 1970 and 49 hours in February -May, 1971. Kenai Moose Research Center. -~eoo --400 JUlY·AU6 (qq IIRS) -~2Do I I ~-t-,00 F'£S·MAY (49 "RS) -~400 -~200 ~OUR .\. ~ •!. Ill 12. t+ lb II!. 2D 2~ OF DAY 53 V> ~ Table 14. Bites per hour by time of day by two male and one female tame moose, 99 hours of observation, July and August 1970, and 49 hours, February-May 1971. Kenai Moose Research Center. Time Period (ADT-hrs.) July-August Totals 4-6 6-8 8-10 10-12 12-14 14-16 16-18 18-20 20-22 (4-22) Number of bites 7124 2403 938 100 3913 4969 5325 2386 123 27,281 Hours observed 14 12 12 6 14 is 14 8 4 99 x bites/hour 508 200 78 17 280 332 380 299 31 237* February-May 8-12 12-14 14-16 16-18 18-20 Totals Number of bites 7166 6356 4569 2459 363 20,913 Hours observed 20.2 13.3 9.5 5.4 1.0 49.4 x bites/hour 355 478 480 455 363 381* * Corrected for number of hourly observations (difference significant P ( .01; t-test). Figure 9. Mean number of bites of natural forage per hour of feeding. 71.95 hours of feeding by two male and one female moose. July and August, 1970 and February -May, 1971. Kenai Moose Research Center. _._1000 500 Dll FEllRUARY-MAY D JULY-AU6USl l·ll I 12·511R5 I 1"'1'11'1! ilrl II •t-6 ~ ~ 10 a2 1~!1-16 \l~ 2 ~ 2."2 ___ l-tC\UR Of= DAY------ 55 Table 15. Mean number of bites taken per hour of feeding by two male and one female tame moose, 42.25 hours of feeding during July and August 1970 and 29.7 hours February-May 1971. Kenai Moose Research Center. Time Period (ADT-hrs.) July-August Totals 4-6 6-8 8-10 10-12 12-14 14-16 16-18 18-20 20-22 (4-22) Bites taken 7124 2403 938 100 3913 4969 5325 2386 123 27,281 Hours fed 12.5 5.0 1.0 0.5 5.5 5.5 7.5 4.5 0.25 42.25 Bites/hour of feeding 571 491 938 200 712 904 711 531 492 646 V1 0'> February-May (Assorted Hours) 8-12 12-14 14-16 16-18 18-20 Totals Bites taken 7166 6009 3708 1235 363 20,913 Hours fed 10.8 8.9 4.9 2.3 0.6 29.7* Bites/hour of feeding 663 675 753 534 622 730 * Some observations over several time periods. that, in some situations, woody twigs are of only minimal importance, contrary to common belief. Physiological studies (eg: Dror et al, 1970) also indicate the importance of variety for proper functioning of the rumen flora. Our tame moose and rumen analysis (described later in this report) studies show that in the 1947 Kenai Burn forbs, lowbush cranberry (Vaccinium vitis-ideae) and lichens are much more heavily used in winter than in summer and that their degree of utilization depends upon snow-determined availability. Although lowbush cranberry becomes more important on depleted range, it is a significant dietary component on normal winter range when available. Lichen seems important only on depleted winter range. Table 15 and Fig. 10 list bites taken of various forage types and species during the 149 hours of feeding observations. Winter observations were divided between the female enclosed in the ten-acre pen of birch- spruce regrowth and two males free-roaming within pen 2 (one square mile). As discussed above, range was considered depleted in the small enclosure, which had supported two animals the previous winter on ca: six acres of land, and had carried three moose throughout the summer. This pen (cf: above) was thus browsed by densities equivalent to approximately 213 moose per square mile (216 x 640) during the preceding winter and by 107 moose-equivalents per square mile during the winter of observations. Despite this, the range was nutritious enough to allow the cow to produce a calf on her second birthday, sired by a 16-month-old bull also living in the small enclosure. Birch leaves made up 56 percent of the mooses' summer diet, forbs 25 percent, sedges, grasses and aquatics 10 percent, willow (Salix ~·) 5 percent and dwarf birch<&· nana) approximately 4 percent (Table 16). Other species, including alder (Alnus crispa), lowbush cranberry and aspen (Populus tremuloides) were included as traces in the diets, as were fungi (Boletus ~.) and soil. Forbs consumed were primarily lupine (Lupinus arcticus)--usually taken in the preflower stage), dwarf dogwood (Cornus canadensis) and wintergreen (Pyrola secunda) in order of preference. On "normal" sera! winter range, (supporting 15 moose/square mile) 76 percent of bites consumed were &. papyrifera woody twig ends. Twenty- one percent of bites were of lowbush cranberry, 3 percent willow, 3 per- cent alder, 1 percent white spruce (Picea glauca) and traces of aspen and highbush cranberry (Viburnum edule). On depleted sera! winter range, only 22 percent of bites taken were woody twig ends of birch. Vaccinium vitis-ideae made up 51 percent of bites, fruiticose lichens (Peltygra ~.) 23 percent, grasses 2 percent, willow 1 percent, foliose lichens (Cladonia ~-) 1 percent and Ribes ~· 1 percent. Forbs, sedges, dwarf birch, aspen, white spruce, fungi and rose (Rosa acicularis) were consumed in trace amounts. Conversion of bite size and numbers to weight of forage consumed is risky due to difficulty in translating visually-estimated bites to collected specimens after a time lapse (cf: Wallmo and Neff, 1970). 57 Figure 10, Plant species consumed by tame moose in summer on normal range and 50 -f-. 25 -~ ;< :.u ~5D -~ ~ -~25 -t- (] 1- ~ ~ 50-- in winter on normal and depleted range. Kenai Moose Research Center, 1970-71. Normal Summer Ranqe --28,423 bites J I I I Normal Hinter Ranqe --10,750 bites I I I I I Depleted Winter Ranne --10 ,lfi3 bites 25------ 58 Table 16. Feeding preferences of two male and one female tame moose on natural forage; 28,423 bites in July and August. Kenai Moose Research Center. Plant Species or Group Vacc. PoEulus B. EaEyrifera Alnus vi tis-tremu1-Total (leaves) Forbs Salix Grasses Sedges Aquatics B. ~ crisEa ideae aides Bites Male -Richard No. Bites 5942 1607 628 70 1 150 709 21 16 9,144 Proportion .65 .18 .07 .01 t .02 .08 t t Male -Walter No. Bites 5277 3847 437 575 253 310 151 1 10,851 Proportion .49 .35 .04 .05 .02 .03 .01 t Both Males No. Bites 11,219 5454 1065 645 254 460 860 22 16 19,957 Proportion .56 .27 .05 .03 .01 .02 .04 t t Female -Raquel No. Bites 4626 1584 448 169 838 485 247 31 8,428 Proportion .55 .19 .OS .02 .10 .06 .04 t All Moose No. Bites 15,845 7038 1513 814 1092 945 1107 22 31 16 28,423 Proportion .56 .25 .OS .03 .04 .03 .04 t t t 0\ 0 Table 16. (cont'd.) Feeding preferences of bvo male and one female moose on natural forage; 21,603 bites in February-Hay. Kenai Hoose Research Center. 10,750 bites by male moose on normal range and 10,163 by a female on depleted range. Normal Depleted Total Plant Species or Group No. Bites Proportion No. Bites Proportion No. Bites Proportion B. £a£Yrifera (stems) 7646 .72 2195 .22 9871 .49 Forbs 14 t 30 t 44 t Salix 335 .03 86 .01 421 .02 Grasses 229 .02 229 .01 Sedges 4 t 4 t Aquatics B. nan a 17 t 17 t Alnus cris£a 364 .03 364 .02 Vaccinium vitis-ideae 2225 .21 5136 .51 7361 .35 Po£ulus tremuloides 18 t 12 t 30 t Lichens Foliose 76 .01 76 t Fruticose 2308 .23 2308 .11 Picea glauca 99 .01 2 t 101 t Viburnum edule 15 t 19 t Fungi 7 t 7 t Ribes sp. 61 .01 61 t Rosa acicularis 2 t 2 t Total Bites 10,750 10,163 20,913 Tables 17 and 18 list distribution of bite sizes on different ranges and during different seasons and mean weights of bites consumed, as estimated by subsequent collections. Combining these data (Table 19) gives estimates of about 4-7 pounds/day wet weight consumption (3 pounds/day dry w~ight) in winter and 42 pounds/day wet weight in summer. The winter estimate is exceedingly low in light of other estimates (eg: Verme, 1970) and our observations that the same tame animals when fed supplementally the pre- ceding winter were each capable of consuming 25-35 pounds pelletized food per day. It thus appears that if previous estimates are accurate, tame moose feeding observations are valuable only in determining proportions of various forages taken in and not for determining biomass relationships. Fig. 11 illustrates seasonal changes in proportions of various foods eaten in relation to snow. Vaccinium was used slightly in late February- early March when snow cover was slight (cf: following sections). During the last half of March and early April, when the Vaccinium availability index (percent of snow plots in which Vaccinium was visible) was lowest, very little was used by moose. From mid-April through May, Vaccinium became· of major importance to moose on both normal and depleted ranges, with Vaccinium and birch ratios (bites) reaching a mean of 15:1 on depleted range and ca:8:1 on normal range. Lichen:birch ratio reached ca:l5:1 on depleted range at this time. Food Habits from Rumen Analysis Tables 20 and 21 summarize plant species and proportions by volume found in rumens of animals shot in seral birch habitat (Game Management Unit 15A) and in climax willow-dwarf birch range (Game Management Unit 7, Juneau Creek). Approximately 80 percent of each sample mean was uniden- tifiable. Of the identified material (Fig. 12) willow comprised 59 percent in the climax range and 39 percent in the 1947 burn. The latter proportion is much higher than expected from this range, where previous estimates (Bishop, 1969) have suggested that willow is less than one percent as abundant as birch in terms of stems per acre. Birch represented 33 percent of the diet in the climax upland range, and 36 percent in the burn; the latter similarly being an under-representation of available birch on the range. Thus, moose are selecting willow over birch where possible, thereby perhaps attaining more variety in their diet (LeResche, 1970). Aspen was present as 8 and 6 percent of the diet in the climax and seral ranges. Lowbush cranberry was a moderately important food item (14 percent by volume) in the seral range, corroborating evidence given by tame moose studies. It was present as a trace in only one of the eight samples from climax range. Forbs (mostly Lupinus ~.) represented five percent of the diets of climax range moose. Rumen protozoa levels on birch and willow and seral ranges and pro- portions of micro-organisms present in rumen liquor are presented in Table 22. Animals from climax willow range had a mean of 283,000 protozoa/ml, statistically equivalent (P) .10; t-test) to those from seral birch range, The higher mean may indicate that the willow range 61 Table 17. Bite size distribution by plant species for two male and one female moose; 45,708 bites; July and August 1970 and February-May 1971. Kenai Moose Research Center. SUMMER Bite Size (no. leaves) ( 5 6-10 11-20 20 Total B. Ea:Qyrifera -no. 6758 6084 2224 789 15,855 - % 42 38 14 5 Forbs -no. 6969 58 10 3 7,040 % 100 t t t Salix spp. -rto. 552 758 176 27 1,513 % -0 36 50 12 2 B. ~ -no. 71 307 361 368 1' 107 -% 6 28 33 33 WINTER Small Medium Large Very Large Total B. 2a2yrifera Normal Range -no. 1413 3285 2458 7696 -% .184 .498 .319 Depleted range -no. 295 760 1151 2206 -% .134 . 345 .523 Total -no. 1708 4045 3609 9902 -% .172 .408 .362 Vaccinium vitis-ideae Normal Range -no. 108 1358 556 177 2199 - % .049 .617 .253 .081 Depleted range -no. 2003 2754 301 135 5163 -% .389 .528 .058 .026 Total -no. 2111 4082 857 312 7362 -% .287 .514 .116 .042 Salix ll· Normal Range -no. .291 23 150 464 - % .628 .050 .324 Depleted Range -no. 50 16 13 79 - % .633 • 202 .165 Total -no. 341 39 .63 543 -% .628 .072 .300 Lichens Normal Range -no. 0 0 0 0 - % Depleted Range -no. 691 1349 346 2386 -% .279 .565 .145 Total -no. 691 1349 346 2386 -% .279 .565 .145 62 Table 18. Wet and dry weights of bites of natural forage taken in February-May 1971. Kenai Moose Research Center. Plant Species Bite Size n Wet Weight (g) Dry Weight (g) % H20 Betula papyri£ era (large) 30 0.66 0.55 17 Betula papyri£ era (medium) 30 0.27 0.22 19 Betula papyrifera (small) 30 0.08 0.05 38 Salix (large) 30 0.78 0.45 42 Salix (medium) 30 0.26 0.15 42 Salix (small) 30 0.07 0.04 57 Vaccinium vitis-ideae (very large) 32 1.13 0.69 39 Vaccinium vitis-ideae (large) 31 0.56 0.35 38 "' Vaccinium vitis-ideae (mediwn) 32 0.32 0.21 35 w Vaccinium vitis-ideae (small) 32 0.14 0.09 36 Comus canadensis (average) 30 0.24 0.12 50 Forbs (large) 30 0.35 0.34 3 Forbs (small) 30 0.04 0.04 0 Grass (large) 30 2.76 2.60 6 Grass (small) 30 0.50 0.49 2 Lichens (large) 30 5.45 1.06 81 Lichens (medium) 30 1.59 0.39 76 Lichens (small) 30 0.67 0.19 72 Birch leaves (July-August) each leaf 500 0.38 ± 144SD Table 19. Estimated consumption of forage by tame moose. Kenai Moose Research Center. Pounds/day wet weight Season Range Birch Forbs Vaccinium Lichens Other Total Winter ''Normal'' 2.6 0.9 0.3 3.8 lbs/day Winter ''Depleted'' 1.0 1.5 4.7 0.2 7.4 lbs/day Summer ''Normal'' 26.7 11.9 42.0 lbs/day Pounds/day dry weight 0\ Winter "Normal" 2.1 0.6 0.2 2.9 lbs/day ~ Winter "Depleted'' 0.8 1.0 1.0 0.2 3.0 lbs/day Figure 11. Ratios of bites Vaccinium vitis-ideae and hites lichen to bites of Betula papyrifera eaten by moose in normal and depleted habitat, February -May 1971. Kenai Moose Research Center. ·::~___ . I ~ ~Aet:INIUM AVAILABIUTY 0.~---------------------------------------------- • ... NORMAL RANt!£ PE PLElED l-tA!»ITAT 0--.-·0 De:PLETE'D AABITAT 2.0 · l Llt\-IEtJ I BETULA) 10 \ \ \ \ ~ ' . \ \ l . ' l . \ \ l ---------DA.TE ------------ 65 Table 20. Food habits of Kenai Peninsula moose as determined by analysis of rumen contents of eight moose collected during the December, 1970 antlerless hunt in Unit lSA (Seral birch range). PERCENT BY VOLUME (of the sample fraction identified) Percent Specimen of Sample PoEulus Vaccinium Dried Mono cots..!/ Forbsl-1 Number Unident. Salix spp. Betula spp. tremuloides vitis-ideae Leaves Others 22794 48 68 8 6 8 8 2 t 22793 91 23 23 35 12 t t 22821 71 70 28 2 t tiS Davis 91 38 58 4 t 22820 65 32 3 t Q\ Q\ #2 Davis 78 17 56 10 16 t t 114 Davis 84 28 24 46 2 t ill Davis 86 31 55 8 6 t Average percent of the eight 78 39 36 6 14 3 t t t Average percent when occurred 78 45 36 12 16 5 t t 2 Percent frequency of occurrence 100 88 100 so 75 88 so 25 so ..!./ Recognizable monocots included: family Gamineae 11 Recognizable forbs included: Pyrola sp. 11 Others included: Rosa sp. and Ledum Table 21. Food habits of Kenai Peninsula moose as determined by analysis of rumen contents of eight moose collected during the December, 1970 antlerless hunt in Juneau Creek, Unit 7 (Climax willow range). PERCENT BY VOLUME (of the sample fraction identified) Specimen Number #3 #6 115 112 Ill ff7 /!4 #8 Average percent of the eight Average percent when occurred Percent frequency of occurrence #3a-'!/ #3~/ ff6a§../ Percent of Sample Unident. Salix spp. 76 85 86 73 86 76 78 85 81 81 100 78 86 72 91 46 60 50 64 50 54 54 59 59 100 90 44 55 Betula spp. 6 43 19 34 27 50 39 46 33 33 100 7 48 37 Populus tremuloides 18 7 t 3 8 38 2 7 ~/ Recognizable monocots included: Carex sp. and family Gamineae 11 Forb species utilized included: Lupinus sp. and unknowns 11 Others included: Ledum and Andromeda ~/ This was a subsample from the same one liter sample as #3. Vaccinium Dried vitis-ideae Leaves t t t 13 2 1 t t t t t t t 62 t t t Monocotsl./ 3 t t 1 38 3 t 21 This was a subsample from the same rumen as #3 and #3a, but from a different one liter sample. ~! This was a subsample from the same rumen as #6. 2/ Forbs- 11 3 9 9 7 5 8 63 Others t t t t t 38 4 4 Figure 12. Identified plant species found in rumens of moose on two types of range. Kenai Peninsula, December 1970. Climax willow -birch ranqe (Juneau Creek; GMU 7) l Seral birch range (Swanson River; GMU 15A) 68 J Table 22. Concentration of protozoans and proportion of sediment material in rumens of moose from climax willow (Game Management Unit 7) and seral birch (Game Management Unit 15A) winter ranges. December 1970. Sample 1 2 3 4 5 6 7 8 Date 2 December 2 December 2 December 2 December 2 December 4 December 5 December Location 7 7 7 7 7 7 7 Sex F F F F F F F Age 5+ yrs. 13+ 18+ 13+ 8+ 14+ Unk. Means for climax willow range 283,000 ± 202,100 SD* 1 2 3 4 5 6 7 8 9 10 16 December 16 December 16 December 16 December 16 December 16 December 15 December 15 December 16 December 16 December 15A lSA lSA 15A 15A lSA 15A lSA lSA 15A F M F F F F M F Unk. Unk. 4+ Calf 7+ 2+ 2+ 12+ Unk. Calf Unk. Unk. Means for sera! birch range 239,000 ± 89,000 SD* * Neither difference significant (P ( .10; t-test). Protozoa/ml 40,000 420,000 130,000 280,000 400,000 140,000 570,000 460,000 240,000 290,000 140,000 230,000 160,000 220,000 230,000 230,000 190,000 % Sedimentation (x of 3 aliquots) 28 36 22 28 34 26 32 30 29% ± 1.5 SD* 32 20 28 23 26 28 32 24 29 26 27% ± 1.2 SD* Fat Index Moderate Heavy Moderate ~1oderate Moderate Unknown Unknown Moderate Light Heavy Noderate Heavy Moderate Moderate Light Unknown Unknown was of higher quality (ie: higher in nitrogen), for Klein (1962) suggested a positive correlation between nitrogen content of rumina and protozoa concentration on summer range. Samples of dried digesta are being analyzed to determine if this is so. The present values for protozoa concentrations are 4-9 percent of values Klein (1962) reported for deer (Odocoileus hemionus sitkensis) in summer. This is logical, for winter range and browse forage are lower in nitrogen than summer forage species and leaves. Proportions of microorganisms in centrifuged factions of rumen liquor were not significantly higher (P ) .10; t-test) in rumens from the upland climax willow range. Chemical Composition of Forage Plants No results are available on this portion of the job. Snow Studies Minor snow studies initiated in February are summarized in Table 23. The studies only touch on this factor that becomes very important to large mammals in many areas (Formozo, 1946; Des Meules, 1964). The primary purpose of snow plots was to determine availability of Vaccinium vitis-ideae to moose during winter. Fig. 13 shows that Vaccinium remained visible in sedge habitat throughout the sampling period and was not visible for only 46 days at most in other habit types. Vaccinium was visible only in sedge type for only 28 days, and at all other times was visible in at least three habitats. During most of the rest of the winter snow was not deep enough to prevent crater-digging moose from feeding on Vaccinium, maximum depth being only 60 centimeters and this persisting for less than 12 days. Although snow cover was associated with consumption of Vaccinium and lichens (cf: Fig. 13) it was only a very minor factor in food availablilty for only a very short time during the winter. In an average winter on the 1947 Kenai burn, ground plants are almost continuously available to moose, especially around the bases of young spruce trees and beneath fallen timbers. Miscellaneous Observations Miller and Parker (1968) reported finding placental tissue in rumens of eight maternal barren-ground caribou (Rangifer rangifer). On 22 May 1970 a piece of tubular tissue approximately 20 centimeters x 1 centimeter was observed hanging from the anus of moose R-70-4, trapped and immobilized in pen 2. The animals's vulva was flaccid from recent parturition and her calf was later observed with her in the pen. RECOMMENDATIONS 1. Moose populations on the northern Kenai Peninsula as represented by penned animals seem to be stabilized or possibly increasing very slightly. 70 Table 23. Snow depth (em.) in each of seven habitat types. Kenai Moose Research Center. 1971. 24 February 4 March 12 March 24 March 31 March 13 April 20 April Mature hardwoods (dense) 8* 27 33 24 22 13* 13* Mature hardwoods (thin) 19 38 47 46 39* 30* 18* Sedge 13* 33* 41* 27* 33* 22* 13* Spruce regrowth 19* 39 44 44* 29* 15* 14* Birch-spruce (thin) 20* 43 51 39 38 33* 22* ....... Birch-spruce (dense) 28* 51 60 47* 36* 26* 23* 1-' Spruce-ledum 28* 48 56 43 41* 36* 28* * Vaccinium vitis-ideae visible. Figure 13. Vaccinium vitis-ideae availability by habitat type. During spring, 1971. Kenai Moose Research Center. HABITAT TYPE Mature Hardwoods (dense) Mature Hardwoods (thin) ..J Spruce Regrowth Birch-Spruce Regrowth (thin) Birch -Spruce Regrovtth (dense) Spruce -Ledum F"EB APR.tL Vaccinium v-i visible Vaccinium v-i not visible 72 Long growth period, great seasonal fluctuations in weight, relatively low neonate survival and high mortality before 24 months of age all suggest a near maximum density of moose is present. This density is likely in excess of 15 animals per square mile. Therefore moose in the 1947 burn should not be managed for increased herd size. 2. Blood sera should be collected from as many weak/moribund animals as possible and from animals maintained on low-quality rations to substantiate suspected nutritional influence on blood parameters. 3. Broken stems per unit area should be further investigated as an indicator of habitat use by moose. 4. 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Young, V. A. & G. F. Payne. 1948. Utilization of "key" browse species in relation to proper grazing practices in cutover Western white pinelands in northern Idaho. Jour. Forestry. 46:35-40. Young, J. w., E. 0. Otchere, A. Trenkle & N. L. Jacobson. 1970. Effect of age on glucose, reducing sugars and plasma insulin in blood of milk-fed calves. Jour. Nutr. 100:11:1267-73. PREPARED BY: Robert E. LeResche James L. Davis Game Biologists APPROVED BY: 88 State: Cooperators: Project No. : Job No.: JOB PROGRESS REPORT (RESEARCH) Alaska Alaska Department of Fish and Game; U. S. Bureau of Sport Fisheries and Wildlife; Kenai National Moose Range; Alaska Cooperative Wildlife Research Unit, W-17-3 Project Title: Moose Investigations Job Title: Moose Behavior Period Covered: July 1, 1970 through June 30, 1971 SUMMARY Five gravid cow moose were radio-collared in May within the Moose Research Center pens. Three other cows and one yearling bull were similarly marked from June through March. The animals were followed to study when and why early calf mortality occurs and to determine patterns of use within the four one-square-mile enclosures. No natural mortality of followed calves occurred after parturition during this study period. Home ranges varied from 94 to 299 acres, or only 15 percent to 47 percent of the available areas within each one-square-mile pen. Overlap of home ranges was considerable (36-47 percent) and activity centers were close to one another, indicating nonrandom habitat use within the pens. Birch-spruce habitat types were used most frequently throughout the year. No hourly preference for habitat types was demonstrated. Distances moved were lowest during late summer and in January and February and greatest in August, November and December. Associations with other moose were highest during rut and in February, and lowest in December, January, and just before parturition. Nonpregnant cows showed reduced sociality during the precalving period similarly to pregnant animals. A student project concerning maternal behavior and the cow-calf bond (Stringham and Lent, 1971) was carried out at the Center during part of the reporting period. i SUMMARY •• BACKGROUND OBJECTIVES PROCEDURES FINDINGS . , Calf Mortality. CONTENTS Home Ranges and Activity Centers. Mobility .... Habitat Use . . Associations of Moose RECOMMENDATIONS. LITERATURE CITED . . BACKGROUND . Page No. . i 1 2 2 6 6 6 10 14 14 19 19 Kenai Peninsula moose calf-cow ratios obtained shortly after calving have consistently been much lower than expected when compared to winter pregnancy data. Pregnancy rates of less than 90 percent in winter have never been recorded yet June calf-cow ratios are commonly less than 40:100 (LeResche, 1970; LeResche and Davis, this report). Most counts made several weeks after calving show substantially lower calf-cow ratios than those made immediately after calving (LeResche, 1968; Kenai NMR Files). These data show the need for determining when and why calves are lost. Past research showed calf mortality was greatest within several weeks post-partum (LeResche, 1968). This research depended upon chance observations of marked calves. Assured relocation of cows by radio- telemetry would increase the chances of rapidly locating still-born calves or recently dead calves so that an autopsy could be diagnostic. Past studies at the Moose Research Center showed a need to determine if patterns of moose use are random within the enclosures. Pellet group and forage production and utilization plots were originally located randomly within habitat types. Analyses of data from these studies suggested other than random use of habitat types (LeResche, 1970). Movements of radio- collared moose could be recorded to test patterns of use. The need to capture and recapture moose for serial blood and milk sampling and measuring and weighing led to the design and construction of traps within the enclosures. Radio locating moose and immobilizing them with cap-chur equipment was one possibility for obtaining serial data if the moose were not trapped at desired intervals. So many studies have suggested the importance of population density and social behavior to productivity (cf: review in Watson and Moss, 1970) 1 that such phenomena have become accepted by some as generally applicable to vertebrates. Still, the most definitive studies have concerned small mammals (eg: Southwick, 1955; Chitty, 1960) and avian species (eg: Lack, 1964, 1966), and any behavioral limitation of large ungulate populations (eg: Wynne-Edwards, 1962; Buechner, 1961) remains conjectural. OBJECTIVES Primary: To locate parturient cows during the calving period and post-partum cows during succeeding weeks to ascertain time and cause of early calf mortality. Ancillary: To determine patterns of moose use within the enclosures. To locate dead animals soon enough for post-mortem autopsy to be diagnostic. To locate animals to obtain needed serial data. To investigate the develop- ment of behavior of moose calves. PROCEDURES Eight cow moose were marked with radios in the 30 mhz range and located daily during late May and early June and at irregular intervals of about one week thereafter. They were located from the ground with a hand-held directional receiver (D 11/m). Transmitters and receiver were obtained from Boyd's Hobby Shop, Tumwater, Washington. All ancillary objectives: Relocating of all radio-collared moose was continued at irregular intervals (approximately one week) until the radios failed or until April, 1971. Each observation was plotted on a 1:10,105 scale vegetation-type map. Habitat type, time of day, activity, date and other moose present were recorded. A student project was initiated during May-August 1970 to investigate the cow-calf behavioral bond and other aspects of calf behavior, as another approach to evaluation of behavioral aspects of productivity. Stephen F. Stringham, a (MS) student at the Alaska Cooperative Wildlife Research Unit, is responsible for the work, which will continue through August 1971. His plan for the study follows: CALF BEHAVIOR AND THE COW-CALF BOND: Stephen F. Stringham Cooperative Wildlife Research Unit University of Alaska OBJECTIVE: To investigate the development of behavior of moose calves. Emphasis will be on social interrelationships and activity patterns. Particular attention will be paid to (a) calving and post-calving periods and, time permitting, to (b) pre-rut and rut periods. 2 I. Social Interrelationships: A. Interactants: 1. cow and calf (or calves) 2. calves of the same cow 3. calf (or calves) and yearling (or yearlings) of the same cow 4. calf-dam pair and other moose 5. calf (when its dam is not involved) and other moose B. Factors to be considered: 1. size and composition (age and sex) of groups inter- acting with calf or calf-dam pair 2. dominance rank--relative to cow--of the other moose 3. spatial orientation of the animals relative to one another a. direction of ears and head b. orientation and posture of body (including "display" characteristics such as erection of the mane) 4. distancing of the moose relative to one another 5. rates of travel toward, away from, or tangential to one another 6. nature of interaction a. maternal b. gregarious c. convergence or avoidance d. play e. "indifference" f. disturbance g. interruption (e.g., interruption of courtship by the dam's calf) h. other 7. which moose initiate(s) interactions 8. frequency, duration, rate and intensity of interaction 3 II. Activity Patterns: A. Activities to be emphasized: 1. social interactions (see above) 2. feeding a. nursing b. browsing or grazing c. drinking d. ruminating 3. resting or sleeping 4. investigation of surroundings 5. play 6. other B. Factors to be considered: 1. time of day, season and year 2. weather 3. duration and frequency of each type of activity (above) 4. location 5. synchronization between activity patterns of calf and dam and between calf-dam pair and other moose with which they are interacting III. Observations during the 1970 field season indicated that onto- genetic changes are both qualitative and quantitative. Close attention will thus be paid to onset of new patterns and trans- formation or extinction of established patterns, and to changes in duration, frequency, rate and intensity throughout the first four months of development. IV. Sample questions typical of those for which answers will be sought in this study: A. In what ways and to what extent does the cow appear to "determine" the behavior of her calf? l, Which activities does the cow initiate; which does she terminate? 4 2. What behavior patterns (e.g., reactions to disturbance) of the cow appear to be mimicked by the calf? 3. Under what circumstances do calves "heel" to their dams? What percentage of the time when traveling with their dams do they "heel"; for how long at a time? What percentage of the time do cows seem to follow their calves, rather than vice versa? B. How does the cow react to "display'' behaviors (e.g., threat postures) by her calf; how do other moose react to them? C. How do calves react to disturbance? Under what circumstances do they drop to the ground or stand and "freeze"; under what circumstances do they flee? D. When a calf is left alone by its dam, how much of the time, and under what circumstances, does it remain bedded down; when it is not bedded down, what else is it doing? E. What differences occur in interactions between male versus female calves and their dams or other moose? PROCEDURES: I. Observations will be made on cow-calf pairs in pens 2 and 4. A. Attention will be concentrated on: 1. The untagged cow with twins in pen 2. 2. R-70-3 in pen 4, and either 3. R-70-1 in pen 4, or 4. R-70-7 in pen 2 II. Though habituation of the moose will be continued, observations will be made while undetected whenever possible. III. Work Schedule: A. Cow-calf pairs will be observed every other week beginning with their first week of life. Since the cow with twins and R-70-3 gave birth a week apart, they will be observed during alternate weeks. The schedule for the third cow to be observed will depend upon when she gives birth. B. The 24 hour day is divided into two periods: 1. A: dawn to 1 p.m. 5 2. B: 1 p.m. to dusk C. Two A and two B periods will be devoted to the cow with twins and R-70-3 every other week; the third cow-calf pair will be observed during one A and one B period on alternate weeks. Thus, six A and six B periods will be spent in the field every two weeks. FINDINGS Of the eight radio-collared cows utilized for the study, four gave birth to single calves, three had none and one died giving birth to twins because of a breech delivery (Table 1). Of the four females producing viable calves, only two cow-calf combinations were followed daily because radio R-70-2 malfunctioned the day after installation and cow R-70-4' abandoned her calf due to human influence (she gave birth while in a trap and was moved from outside the enclosures to within). Observations of the two viable cow-calf combinations indicated that luck would be involved in locating a recently dead calf via the cow unless she remained by the dead calf for many hours. Even within the first two days of the calf's life, one cow and calf moved a minimum of 460 meters (1,380 feet) in heavy cover. This illustrates that even with daily checks, locating a dead calf would be difficult. Home Ranges and Activity Centers Characteristics of home ranges and activity centers were calculated from 175 observations involving six radio collared moose (Table 2). The number of observations upon which the following discussions of home ranges, activity centers, mobility, habitat use and associations are based are presented in Table 3. Many researchers have utilized, and defined differently, home range and activity center concepts in calculating areas of animal activity (Mohr and Stumpf, 1966; Robinette, 1966; Sanderson, 1966; Bayless, 1969; Hawkins and Montgomery, 1969; Rongstad and Tester, 1969; Goddard, 1970, Telfer, 1970; and VanBallenberghe and Peek, 1971). Activity centers were determined by the method of Mohr and Stumpf (1966). This entailed locating a major and minor axis each of which divided the radio location points into two equal groups, while making the sum of the distances of all points from each axis as small as possible. All radio location points were utilized. Activity centers of moose in pens 2 and 4 were much closer together than would be expected by chance (Fig. 1), suggesting nonrandom use patterns by selection of areas of better habitat. Home ranges were calculated after Godfrey (1954) from Mohr and Stumpf (1966). This method consisted of connecting all outside points of observation and calculating the area of the enclosed polygon. 6 Table 1. Identity of radio-collared moose in the Moose Research Center during May 1970 to April 197L Moose Radio Color Installed Quit Pen Sex May 1970 Age. Yrs. Reprod. 19 70 & Remarks R-70-1 Gr. & Blk. 19 May 70 16 June 70 4 F 6 1 calf born 20 May R-70-2 Blk. & Silver 22 May 70 23 May 70 2 F 3 1 calf born 22 May R-70-3 Org. & Yellow 20 May 70 22 Oct. 70 4 F 3 1 calf born 23 May R-70-4 Blu/wh. 23 May 70 Working 2 F ? 1 calf born 23 May deserted R-70-5 Red & Gr. 24 May 70 Working 2 F ? Female died, 2 calves, 1 breech R-70-7 Red & Gr. 4 June 70 Working 2 F 7 R-70-8 Gr, wh, yel. 10 July 70 Working 1 F 2 R-69-3 Gr. & Blk 13 Aug. 69 Working 2 F 6 Richard Bk, yel, wh. 11 Nov. 70 20 Nov. 70 2 M 1 Richard Yel, Red 30 March 70 Working 2 M 1 Table 2. Observations of each of six radio-collared moose by month. Kenai Moose Research Center. 1970-71. Pen 1 Pen 2 Pen 4 ALL PENS R-70-8 R-69-3 R-70-4 R-70-7 Total R-70-1 R-70-3 Total May 1970 4 4 8 9 9 18 26 June 11 10 8 29 9 10 19 48 July 3 3 5 4 12 2 4 6 21 August 4 5 2 4 11 3 3 18 September 3 3 3 3 9 4 4 16 October 2 2 1 2 5 2 2 9 November 3 2 2 3 7 10 December 2 2 2 2 6 8 January 2 1 1 2 4 February 2 3 2 2 7 9 March 2 1 1 2 4 April 1 1 2 2 TOTAL 23 38 32 30 100 20 32 52 175 Table 3. Characteristics of home ranges Research Center. 1970-1971. Home Range Pen Moose (acres) (acres) R-70-8 175 640 R-69-3 299 640 R-70-4 247 640 R-70-7 234 640 (partial) R-70-3 148 640 R-70-1 107 640 (complete) R-70-3 94 640 00 Acres Connnon to Home Ranges of: R-69-3 ) R-70-4 ) 109 acres R-70-7 ) R-69-3 ) R-70-4 ) 172 R-69-3 ) R-70-7 ) 145 R-70-4 ) R-70-7 ) R-70-1 ) R-70-3 ) 53 of radio-collared moose within one-square-mile enclosures. Kenai Moose Percentage Times Dates Located Index of of Pen Located From To HR size Pen 27 22 14 July 70 30 Mar. 71 .59 1 47 40 22 May 70 21 Apr. 71 1.00 2 39 33 24 May 70 30 Mar. 71 .83 2 37 31 5 June 70 14 Apr. 71 0 78 2 23 22 21 May 70 14 July 70 .so 4 17 20 21 May 70 14 July 70 0 36 4 15 32 21 May 70 22 Oct. 70 .31 4 Percent of Individual's Home Range Overlapping with Other Moose: R-69-3:36% R-70-4:44 R-70-7 :47 R-69-3:58 R-70-4:70 R-69-3:49 R-70-4:59 R-70-4: R-70-7: R-70-1:50 R-70-3:56 I Figure 1. Activity centers and home ranges of radio collared moose within the Kenai Moose Research Center. PEN I --R·70·8 ONE MIL£ fEN 2 R·IOCJ·' ----R.·7o·4- ········R·70·7 PEN4 ----. fEN3 ...... I -.- / ( Determining home ranges and activity centers for moose confined within one square mile may be of limited usefulness as home ranges might likely include the entire one mile if enough observations were made over a long period. However, it is interesting that during one year, with a con- siderable number of observations, individuals moved over a relatively small portion of each one square mile enclosure (Table 3; Fig. 1). These home ranges were also considerably smaller than those (2-10 square miles) reported for free-ranging moose by Phillips and Berg (1971). This may be due to the enclosures or the the excellent and diverse habitat present in small areas in the 1947 Kenai Burn. The home range data also suggest that utilization of the pens is probably not random. While none of the three radioed moose in pen 2 had a home range encompassing over 47 percent of the one-square-mile pen, all three of the animals shared a common portion of the pen that amounted to 36, 44 and 47 percent, respectively of their total home ranges (Table 2). Also, activity centers for R-70-4 and R-70-7 were only 190 meters apart and that of R-69-3 was about 600 meters from the others. This may be due partially or wholly to confinement. Two radioed animals in pen 4 utilized only about one-fifth of the pen during a 10-week period, Both animals were caught in a trap in that portion of the pen. Further evidence that penned moose have home ranges of relatively small size is that individually marked moose are often predictably observed by chance in small portions of the pens. Mobility Average distance moved between observations by month (Figs. 2 and 3), average distance moved between all observations, and greatest and least movement between two consecutive observations (Table 4) were computed from the radioed moose relocations. Utility of these data is obviously restricted because of limits on movements placed by the enclosures. However, minimum distances moved and differences in distances moved by month perhaps indicate true movement patterns. Also, the small proportion of the total pen area used by each animal indicates that the pens are not as restrictive as might be expected. Observations were made at irregular intervals, varying from one day to over a week. However, relative monthly average distances between relocations correspond closely for most moose in all pens. The exceptions were that average distances moved: 1) decreased from July to August for all except pen 1, where they increased; 2) all decreased from August to September except for that of the cow in pen 4, which increased slightly (possibly because of calf development); and 3) decreased sharply in all pens from December to February except in pen 1 where it increased from January to February (possibly accounted for by few observations). The most striking inter-pen differences were: 1) the much smaller average distance moved in pen 4. This was expected because the two cows here were the only ones studied that had live calves. Their average distance moved increased from June to July as expected because of increas- ing calf mobility with age but decreased again for August and September. 10 Figure 2. Mean distance between radio collared moose relocation for individuals in Pen 2. M h C 1970 1971 Kenai oo~e Researc enter. -. R.1{J 4 ~ ~q .. ~ \ . . -. --1-I \ ·--R.70· 1 . . \ I \ . -r-4000 I \ -. \ . I \ . . -r-I \ I~ ~ \~ . . . \ I I \ I . . -300£) \ ! . -\ I \ r-- i ~ . I I \- I \ . \\ . \ . --I I \\ - \ f v . \.\ I ' I . r-2/VV\ I \ I ·--. \\ I\ I \\ . I "'I \ . -1-\ \ - i\v 1 . \ ~ • . I \' -r-IMO \ r--. \ / ~\ v 5 6 7 S q 10 II 12. I 2. ----IV\.01\.JT"f-1 ----- 11 -----~~---------·-···------- Figure 3. Mean distance between radio collared moose relocations for all pens occurring within each month. Kenai Moose Research Center. 1970 -1971, MEMJ: PGJ2 (.3MOOSE.) --MEAN: All PENS (6 MCilSE) 11,0 _. _ MEAN:PEN4-(2 ~t) _ .. _MFAN: PEN I .(l MOOSE) <I] ~ ~ I~ ' (/) ~ -~ \ \ (.IJ IX • \ ! 670 I • ·. \ • \\ I • I "\ Ul !500 • (LJ I I ~-~ (J . z I \ . \ • ~ . I / \ . ' VJ 330 • . • • -0 .,..., -.. 2 l6 ~ 160 MAV J J A ~ 0 N D J MONTl-4 12 F' Table 4. Greatest, least and mean distances moved between radio relocations of moose. Kenai Moose Research Center. 1970-71. Distance between relocating J20ints ~m~ Moose Pen Greatest Least Mean R-70-8 1 1560 25 607 R-70-7 2 1840 184 688 R-70-4 2 2160 44 680 R-69-3 2 70 813 All 2 99 724 R-70-3 4 1883 53 486 R-70-1 4 1250 33 386 All 4 1570 43 436 13 The consistently smaller monthly average distance moved for pen 1 compared to pen 2 possibly occurred because of having data from only one individual in pen 1. That animal may have been a more sedentary individual or perhaps restricted movement reflected the habitat juxtaposition of cover types. Other noticeable changes in average distance moved per month were: 1) the decrease from May to June for all animals without calves; 2) the increase from June to July and subsequent decrease to August (the August decrease coincided with a shift to mature hardwood type suggesting pre- rut behavior alteration or perhaps a seasonal preference for the type); and 3) the low average distance between movements during the months of September and October, (associated with the breeding season) the sub- sequent increase after the rut, and the decrease as winter progressed. Habitat Use Considering observations of radioed moose as an index of habitat use, birch-spruce regrowth types were by far the most heavily utilized vegeta- tional type throughout the year (Fig. 4). On a seasonal basis this type received over 50 percent of the total use during all months except August and September when it received 22 and 33 percent of total use, respectively. During these two months mature hardwood was utilized at rates of 61 and 40 percent, respectively. The only other habitat types receiving much utilization were sedge and spruce-ledum from May through October. Utilization of sedge and spruce-ledum types occurred (primarily by R-70-1 and R-70-3 and their calves) from May through September. Several observa- tions in other pens occurred in these types during this period also. Spruce regrowth and mature spruce type utilization was negligible during most months. Percentages of total observations of each individual occurring in each habitat type were quite consistent with the exception of pen 4 individuals (Figs. 5 and 6). 'Both animals here were observed more often in sedge, spruce-ledum, and hardwood types, and less frequently in birch- spruce type than were other moose. This likely occurred because the pen 4 animals had calves. Associations of Moose The association of each radioed moose with all other moose with which it was observed was quantitatively expressed (Table 5). The method used for computing was that which Knight (1970) presented as: "Formula for the coefficient of association: 2 ab a + b Where a is the number of times animal A was observed through- out the season, b is the number of times animal B was observed throughout the season, and ab is the number of times that animals A and B were observed together throughout the season. 14 Figure 4. Percent of total monthly observations of all radio collared moose by habitat type. Kenai Moose Research Center. 1970-1971. llJO 70 oO 20 Spruce-Led urn n=12 Sedge n=16 Mature Hardwoods n=41 10 Mature Spruce n=4 Spruce Regrowth n=l Birch-Spruce Regrowth n=86 MAY J a N D J 26 44 B to 6 3 15 Figure 5. Percent of total occurring in each observation of radio collared moose in Pen 4 by habitat type. Kenai Moose Research Center. 1970 -1971. 60 R 70-1 with one calf -• •-• Mean 50 - - - R 70-3 with one calf /\_I CALF I \ I \ I \ I (' . . • I . <I) I / 2 0 • -1! 1- ~ I: a.! tu 20 \ ,, ~ • '\. ~· (- ~ . f ' ,\. 10 ~ "./ SE06£' SPRUe£· AAR.l»U!tJ 61RCM SPI?Ut£· Sf'RlJC£" LEDUM SPRllC£ RE'610NAl MATUR.£' -------HAB\TA.\ TVf'£ -------- 16 Figure 6. Percent of total observations of radio collared moose in each pen by habitat type. Kenai Moose Research Center. 1970-1971. N: SED6E Ito SPRUC.E' HABMia)£) BIRO- L£Dllt.J\ , s~E' 12. 41 Bil ------~ABITA\ \'fPc _____ _ 17 ...... 00 Table 5. Coefficients of association for moose within three one-square-mile enclosures. Kenai Moose Research Center. 1970-71. Moose 43 M 40 F 6 F Mean R-70-8 F .28 .07 .07 .14 1 F 36 M R-70-7 9 F R-69-3 F .08 .18 .17 .12 R-70-7 F .16 .23 .21 R-70-4 F .33 .24 .16 All pen 2 .02 .25 .20 .16 21 M (before calving) Calf R-70-1 F .13 .80 R-70-3 F 1.0 Pen 2 means: males= .13 females = .14 Association With R-70-4 45 M Rich Walt R-69-3 Unmarked Mean .09 .04 .14 .08 .11 .24 .18 .og .06 .17 .06 .16 .12 .OS .05 .09 .06 .16 .11 .09 .06 .13 .04 Using this formula, a value of 1 would indicate perfect associa- tion or the probability that A and B would occur together all of the time." As shown in Table 5, there was no strong association between any animals other than the two cows and their calves in pen 4. Even these associations were followed only through the first four months of the calves' lives. An explanation for the less than 1.0 association between R-70-1 and her calf is that the calf was not always observed when the cow was located in heavy cover. Even during her calf's first week of life, R-70-3 was observed over 300 meters from it, illustrating that the animals may not be observed together even while closely associated behaviorally. The nearest day to a parturition date that either cow was observed associated with another adult animal was when R-70-1 was observed with male #21 on May 23, five days before her calf was born on May 28. There was no significant difference in degree of association between individuals within the various pens, individuals of one pen compared with those in another, or between sexes. The highest degree of association occurred during the rut and during February (Fig. 7). These were also the times of most restricted movements. The February high was unexpected in light of the lowest monthly associa- tions of the year occurring during December, January, and March. Even though the four cows in pens 1 and 2 had no calves, they all showed a lowered degree of association during the pre-calving period of May, similar to the gravid cows in pen 4. RECOMMENDATIONS 1. Radio-telemetry of parturient cows is an unpromising way to study early calf mortality. If this type of study is to be pursued, a method should be found for radio-tracking calves themselves or for enclosing cows into areas small enough to allow location of calves but large and ecologically complete enough to support a cow and her calf. 2. One-square-mile pens appear large enough to enclose moose at normal densities without unduly restricting movements and home ranges. LITERATURE CITED Bayless, S. R. 1969. Winter food habits, range use, and home range of antelope in Montana. J. Wildl. Mgmt. 33(3) :538-551. Buechner, H. K. 1961. Territorial behavior in Uganda. Kob. Science 133:698-99. 19 Figure 7. -100 -50 -70 -60 -50 -40 -10 Percentage of total monthly observations of radioed moose in Pens 1 & 2 that involved associations with other animals. Kenai Moose Research 1970-71. 8Y SELF .r A s 0 D J F M 2.1 IS 16 q 10 4 q N.UM&ER O.BS. 20 Chitty, D. 1960. Population processes in the vole and their relevance to general theory. Can. J. Zool. 38:99-113. Goddard, J. 1970. Movements of moose in a heavily hunted area of Ontario. J. Wildl. Mgmt. 34(2):439-445. Godfrey, G. K. 1954. Tracking field voles (Microtis agrestis) with a Geiger-Miller counter. Ecology 35(1):5-10. Hawkins, R. E. and G. G. Montgomery. 1969. deer as determined by telemetry. Kenai NMR Files. (unpublished) Movements of translocated J. Wildl. Mgmt. 33(1):196-203. Knight, R. R. 1970. The Sun River elk herd. Wildl. Monog. 23:66 p. Lack, D. 1964. A long-term study of the Great Tit. J. Anim. Ecol. 159-73. Lack, D. 1966. Population Studies of Birds. Oxford. 341 p. LeResche, R. E. 1968. Spring-fall calf mortality in an Alaska moose population. J. Wildl. Mgmt. 32(4):953-956. LeResche, R. E. 1970. Moose report. Annual Project Segment Report. Vol. XI. W-17-2. LeResche, R. E. and J. L. Davis. 1971. Moose report. Annual Project Segment Report. Vol. XII. W-17-3. (This report). Mohr, C. 0. and W. A. Stumpf. 1966. Comparison of methods for calculat- ing areas of animal activity. 30(2):293-303. Phillips, R. L. and W. E. Berg. 1971. Home range and habitat use patterns of moose in northwestern Minnesota. 7th N. Am. Moose Conf. (abstract). Robinette, W. L. 1966. Mule deer home range and dispersal in Utah. 30(2):335-348. Rongstad, 0. J. and J. R. Tester. 1969. Movements and habitat use of white-tailed deer in Minnesota. J. Wildl. Mgmt. 33(2):366-379. Sanderson, G. C. 1966. The study of mammal movements--a review. J. Wildl. Mgmt. 30(1):215-235. Southwick, C. H. 1955. The population dynamics of confined house mice supplied with unlimited food. Ecology 36:212-25. Stringham, S. and P. C. Lent. 1971. Calf behavior and the cow-calf bond in moose. Work Plan Seg. Rept. W-17-3. R-197. 22 p. (Xerox) 21 Telfer, E. S. 1970. Winter habitat selection by moose and white-tailed deer. J. Wild!. Mgmt. 134(3):553-558. VanBallenberghe, J. and J. M. Peek. 1971. moose in northeastern Minnesota. Radiotelemetry studies of J. Wild!. Mgmt. 35(1):63-70. Watson, A. and R. Moss. 1970. Dominance, spacing behavior and agression in relation to population limitation in vertebrates. In Watson, A. (ed.) Animal Populations in Relation to their Food Resources. (Oxford). p. 167-220. Wynne-Edwards, V. C. behavior. PREPARED BY: Robert E. LeResche James L. Davis Game Biologists 1962. Animal dispersion in relation to social Edinburgh & London. APPROVED BY: of Game 22 State: Cooperators: Project No. : Job No.: JOB PROGRESS REPORT (RESEARCH) Alaska Alaska Department of Fish and Game, U. S. Bureau of Sport Fisheries and Wildlife (Kenai National Moose Range) W-17-3 Project Title: Moose Investigations Job Title: Development and Testing of New Techniques Period Covered: July 1, 1970 through June 30, 1971 SUMMARY Techniques of aerial censusing, pellet-count censusing, immobilizing, radio tracking, freeze-branding and otherwise marking moose were tested in the Moose Research Center enclosures and nearby. Experienced observers saw a mean of 69 percent of moose flown over in excellent counting conditions, when flying 15 minutes over each one- square mile. They saw significantly (P ( .02; t-test) more moose than experienced observers in good conditions, (61 percent), and experienced observers in poor conditions (40 percent). In all conditions, experienced observers saw more than did inexperienced observers (43 percent, 44 percent and 19 percent, respectively for excellent, good and poor conditions). Observers with past experience but little or no current experience saw a proportion of moose (46 percent) equivalent (P ) .10; t-test) to that seen by inexperienced observers and significantly lower (P ( .01; t-test) than that seen by experienced/current observers (excellent conditions). No significant (P ) .10; x2 ) difference in ability to see moose was detected in relation to time of day, although highest proportions were seen between 1000-1200 hours and after 1400 hours, in March. Counting of pellet groups within 159 plots cleared of pellets in 1970 suggested a defecation rate of 32 groups per day, or 0.59 ± 0.90 groups per plot deposited in one year. M-99 Etorphine and succinylcholine chloride were drugs of choice for moose. Addition of hyaluronidase to injected drugs increased absorption rate and apparently stabilized drugs' effect on different individuals. Effects of the drugs varied seasonally. Other drugs were less useful with moose. Freeze-branding experiments were unsuccessful on two moose. An easily read collar-pendant marker was placed on 135 adult moose. i SUMMARY •. BACKGROUND. . OBJECTIVES. . PROCEDURES. . FINDINGS .. Aerial Census Evaluation Pellet-count Census. Immobilizing Drugs Radio-Telemetry •.• Freeze-Branding·. . . Other Marking Methods .. RECOMMENDATIONS . LITERATURE CITED. . . . . . CONTENTS BACKGROUND Page No. i 1 2 3 5 5 12 12 16 16 16 16 17 Moose research and management require methods of estimating numbers and of handling, marking and following animals. These techniques necessarily vary with species, location and nature of the management/ research problem. The Moose Research Center, with known numbers of con- fined animals, provides a unique test-ground for numbers-related techniques and for methods and equipment whose effectiveness can be learned only by relocation of animals. Aerial censusing is at present the only practical method of estimat- ing moose numbers in most of Alaska (cf: Rausch and Bratlie, 1965; Rausch and Bishop, 1968; Bishop, 1969; Bergerud and Manuel, 1969), but determina- tion of the extent to which this method underestimates numbers has been a major problem when absolute numbers are sought. Benson (1966), Goddard (1966) and Bergerud (1968) have reviewed aerial censusing techniques. LeResche (1970) reported that moose aerial census results depended upon observers and counting conditions and that fewer than 70 percent of moose present were seen by experienced observers flying with good count- ing conditions. Siniff and Skoog (1964) developed a random stratified, quadrat sampling method for caribou (Rangifer tarandus), but even in intensively counted quadrats, some animals were missed. Evans et al (1966) used a similar technique on moose. The presence of four one-square-mile pens with known numbers of moose provided an opportunity to test the accuracy of aerial censusing and to test the value of previous experience in aerial counting. Pellet-count census techniques have been used for various species of big game animals since the 1930's (cf: Bennett~ al, 1940; Rasmussen and Dovan, 1943; Bowden et al, 1969). Several studies have been done 1 with penned ungulates (summary in Neff, 1969) and others have used the technique in intensive studies of habitat use (DesMeules, 1962). The known numbers of animals enclosed in the Moose Research Center also pro- vided opportunity to test this population-estimation technique for moose. Immobilization of big game animals by drugs has progressed rapidly in a very few years, with different drugs indicated for different species (cf: Harthoorn, 1964, 1968; Miller, 1968; Houston, 1969). An ideal immobilizing drug should have 1) short induction time, 2) wide tolerance range, 3) rapid reversibility, 4) no lasting or cumulative side effects and 5) should leave meat consumable by a subsequent hunter. To find such a drug for a given animal, the logical approach is to try agents as they become available, preferably on recoverable animals. Succinylcholine chloride (Bergerud ~ al, 1964) has been the most often used drug for immobilizing moose in Alaska primarily because it alone fulfills condition (5) above. The drug is less than ideal because it has a very narrow range of safe dosage (20 percent or less--LeResche, 1970), is irreversible and leaves moose with no muscle control to resist drowning in wet places. A recent report (Fuyita, 1970) has documented a delayed hypersensitivity reaction to the drug in a human subject. Pienaar (1968a, b) has reported in detail on effects of various thebaine derivatives (eg: M-99 Etorphine) when used alone and in com- bination with tranquilizers and parasympatholytic agents. These neuroleptic- analgesic mixtures proved ideal because of their reversibility, wide safety range (230 percent for M-99 administered to moose reported by LeResche, 1970) and favorable therapeutic index. Houston (1969) and LeResche (1970) reported on M-99's effectiveness in handling moose. Migration and other behavioral studies of big game animals may be accomplished by marking many animals and searching for them. Recently, radio-tracking devices have come into vogue for continuous location of animals (cf: Slater, 1963 for one review). In programs requiring regular relocation of moose for sampling blood or other specimens, radio-tracking gear is invaluable to insure timely recapture. Methods of marking moose for subsequent identification have stressed assorted collars, pendants and earflags. In work with domestic animals, freeze branding (Farrell et al, 1969; Kambitsch et al, 1969) has recently come into greater use. The method involves killing pigment-producing cells in hair follicles by freezing and produces white hair in the pattern of the brand applied. OBJECTIVES To develop and for test techniques for: aerial censusing, pellet- count censusing, immobilizing radio-tracking and marking of moose. 2 PROCEDURES On January 26-February 4, 1970, three helicopter counts and 19 counts by PA18-150 supercubs were made of moose in the four Moose Research Center pens. Thirty-three additional supercub counts were made on 6-9 March, 1971. Observers were instructed to direct pilots how to fly the survey and were allowed 15 minutes to count each square mile. Pilots did not participate in locating moose and observers recorded each moose seen. Observers could direct pilots to circle over one small area, to fly tran- sects, dive, etc. Conditions were good-excellent, with snow cover at least adequate, for 15 counts in 1970 and poor for 4 that year. Conditions were excellent, with complete snow cover, during all 1971 counts. Time of day, pilot and previous moose-counting experience of the observer were noted for each observer and the total number of moose seen in each pen was recorded. In spring 1970 (LeResche, 1970) 728 permanent 8' x 24' browse- utilization plots.were used as pellet-count plots. Fecal groups in each plot were classified as "winter" (pellets) or "summer" (non-pelletized) , counted and cleared from the plot. Data analysis was by habitat type and an estimate was made of required sampling intensity. Pellet groups in two pens were separated into "new" (the preceding winter) and "old" by guess, and endurance of pellet groups was estimated from these data. Habitat use index was calculated from numbers of pellet-groups. On 2-4 June 1971, newly deposited pellet-groups were counted on 159 plots in Pen 1. These plots had been cleared of pellets in spring, 1970. M-99 Etorphine and M-50-50 Diprenorphine were tested on 28 trapped animals sine€ October 1970. An additional 116 animals were immobilized using succinylcholine chloride. Hyaluronidase was used to speed absorp- tion of the drugs. One animal each was handled with sernalyn and sparine, "Bay-Va" (Bauer-Werke, Belgium), succinylcholine chloride and sodium pentabarbitol, and succinylcholine chloride and "tranvet". Radio transmitters in the 30 mhz range were used to periodically relocate 10 moose within the pens (cf: Job 1.2R). On 10-11 November 1970, two tame 17 month-old male moose (Walter and Richard) were freeze-branded. Hair was shaved off with electric animal clippers and cold copper branding "irons" were applied to the skin. Numbers were 6" high and approximately 1" wide (L & H Mfg. Co., Mandan, S.D. 58554). Irons were cooled with a mixture of acetone and dry ice and applied for various lengths of time (Table 1). Thirteen x eighteen em (5 x 7") pendants with 8 em (3") high by 8 mm (5/16") wide letters and numerals (eg: "Al-AlOO"; "Cl-ClOO'') were suspended from collars under the necks of 135 moose immobilized from helicopters in the Moose River Flats. Pendants were of laminated plastic, red outside- 3 Table 1. Experimental applications of freeze-brands to moose. Kenai Moose Research Center, November 1970. Moose Site Brand Number Time Applied Walter Right Flank 3 40 seconds Right Rump 2 20 seconds Left Shoulder 9 50 seconds Left Rump 8 60 seconds Richard Left Shoulder 0 60 seconds Left Flank 5 20 seconds Left Rump 4 40 seconds Figure 1. Canvas-webbing collars placed on moose, with colored plastic facing material for individual identification. Lays on moose's dorsal side t A A A A E D c B B B B A: Red, orange, pink, yellow. B: Red, orange, pink, yellow. C: Green, brown, black, silver, yellow. D: Green, brown, black, silver, yellow. E: Green, brown, black, silver, yellow. 4 white inside. Numbers were routed out of both sides so that they appeared as white numbers on a red background. Seventy pendants were hung perpen- dicular to the mooses' longitudinal axis (ie: facing forward) and 65 were hung facing sideways. Attempts were made to read pendants from supercub aircraft. In May 1971, 63 canvas-webbing collars each with a unique stripe combination (Fig. 1) were placed on free-ranging moose. These collars also carried pendants (cf: above). Attempts were made to determine animal identity from supercubs using these color combinations. FINDINGS Aerial Census Evaluation Table 2 presents new data of number of moose seen by individual observer in 1971. Similar data for the 1970 counts were presented in Table 25 of LeResche (1970). Table 3 summarizes mean results by year, conditions and counters' experience. Results with experienced observers in good conditions (61 percent moose seen in 1970; 69 percent in 1971) differ significantly (P ( .02; t-test) between 1970-71, but inexperienced observers in good conditions performed equivalently (44 percent vs 43 percent; P) .10; t-test) both years. The difference among experienced observers is probably related to better snow cover in 1971 (see below). However, since inexperienced observers in good conditions did not vary between the two years, they are lumped in the following analyses. Proportion of moose seen during aerial surveys depended upon: observer, snow cover (counting conditions), pilot, time of day, aircraft and (to the extent we could determine) terrain. Effect of Observer Skill Under all conditions tested, experienced observers (those who had counted game often before and had done so recently) saw a significantly greater (P ( . 01; t-test) proportion of moose flown over than did inexperienced observers. In excellent conditions (1971), experienced observers (n = 12) saw 69 percent of the moose they flew over. Inexperienced observers (n ~ 10) saw but 43 percent. In good conditions (1970), experienced observers (n ~ 4) saw 61 percent, inexperienced (n = 11) saw 44 percent. In poor conditions, three experienced observers saw 40 percent. One observer with no experience saw 19 percent. Variation among observers with current experience was great (ranges: 45 percent-80 percent, 49 percent -76 percent and 35 percent -44 percent in excellent, good and poor conditions, respectively) as it was among inexperienced observers (ranges; 33 percent -50 percent; 27 percent -61 percent in excellent and good conditions, respectively). 5 Table 2. Results of moose counting experiment by individual counter: 6-9 March 19 71. Pen (number present) Rank in Inexperienced Time Pilot 1(12) 2(16) 3(12) 4(15) Tota1(55) Percent Group 1 11 am II 8 8 5 8 29 53% 1 2 1 pm I 6 8 3 10 27 49% 2 3 10 am I 3 13 4 7 27 49% 2 4 10 am I 7 10 6 3 26 47% 3 5 9 am II 6 7 7 5 25 45% 4 6 12 noon I 7 4 6 5 22 40% 5 7 llam II 5 6 5 5 21 38% 6 8 12 noon II 4 9 4 4 19 35% 7 9 1 pm II 6 5 4 4 19 35% 7 10 1 pm I 5 3 3 7 18 33% 8 11 10 am II 4 )counted only Pen 1 12 2 pm I 7) Experienced-but-not-current • 13 9am I 6 8 9 9 32 58% 1 14 3 pm I 5 13 8 6 32 58% 1 15 3 pm I 8 10 6 7 31 56% 2 16 4 pm II 7 10 6 7 30 55% 3 17 7am I 6 6 5 7 24 44% 4 18 12 noon I 7 5 6 6 24 44% 4 19 8 am II 5 7 3 6 21 38% 5 20 9 am II 4 7 3 3 17 31% 6 21 12 noon I 2 4 4 6 16 29% 7 Ex~erienced-Current 22 11 am I 12 14 8 10 44 80% 1 23 2 pm II 10 8 9 15 42 76% 2 24 3 pm I 11 11 8 12 42 76% 2 25 3 pm I 6 13 6 15 40 73% 3 26 4 pm II 8 9 9 14 40 73% 3 27 1 pm I 9 13 7 10 39 71% 4 28 3 pm II 11 10 7 10 38 69% 5 29 12 noon II 12 10 4 11 37 67% 6 30 10 am I 9 9 7 12 37 67% 6 31 1 pm I 6 10 8 11 35 64% 7 32 9 am I 9 7 6 11 33 60% 8 33 3 pm II 8 6 4 7 25 45% 9 Table 3. Summary of mean proportions of moose observed in four one-square mile enclosures by observer, weather conditions and aircraft. January-February 1970 and March 1971. (Each square-mile searched 15 minutes.) Observer (n) 1970 Pilots (2) plus observers Experienced (4) Experienced (3) Inexperienced (11) Inexperienced (1) Experienced* (3) * 2 Observers 1971 Inexperienced (12) Experienced-but-not- current (9) Experienced-current(l2) All (1971) (33) Conditions Good Good Poor Good Poor Excellent Excellent Excellent Excellent Aircraft PA 18-150 Supercub PA 18-150 Supercub PA 18-150 Supercub PA 18-150 Supercub PA 18-150 Super cub BG 4A Helicopter PA 18-150 Supercub PA 18-150 Supercub PA 18-150 Super cub I(7) 1.00 .82 .43 .57 .14 .93 .47 .46 .77 .58 Pen (n) Total {n) II(l2) III(l2) IV(l8) .75 .71 .64 .73(49) .56 .67 .53 .61(49) .33 .44 0 39 .40(48) .46 .45 .37 .44(49) .25 .25 .12 .19(48) 0 78 .58 .80 .75(48) .46 .37 .39 .43 .48 .46 .42 .46 .63 .58 .77 .69 .53 .48 .54 .53 Currency of experience was important. Nine observers who had regularly counted animals from aircraft at one time but had not done so within one to several years of the experiment saw only 46 percent of the moose flown over (range 29 percent-58 percent) in excellent conditions. This proportion is statistically equivalent (P ) .10; t-tes t) to that of inexperienced observers and significantly lower (P ( .01; t-test) than that of experienced/current observers. Effect of Counting Conditions As suggested above, counting conditions significantly aff~cted proportion of moose seen during aerial counts. As used here "conditions" roughly means "snow cover". Other factors such as turbulence and light conditions are important also. However, turbulence is usually minimal when flights are made and lighting depends primarily upon time of day (cf: below). "Excellent" counting conditions as used here indicate the 40-45 em (16-18") of snow present on 6-9 March 1971 (cf: Job l.lR). This amount of snow covered essentially all dark spots on the ground and clung to some small spruce trees. Good conditions (1970) were approximately 20-25 em (8-10") snow cover, with a few bare patches and no spruce snow-covered. Poor con- ditions (1970) included only patches of snow on the ground. Experienced observers saw significantly (P ( • 01; t-test) more (69 percent) moose in excellent conditions than in good (61 percent) and significantly (P ( . 01; t-tes t) more in good conditions than in poor con- ditions (40 percent). Inexperienced observers saw statistically equivalent proportions ( ) .10; t-test) of moose in excellent (43 percent) and good conditions (44 percent) indicating variation in snow conditions (when some snow cover does exist) is important only to experienced observers. Only one inexperienced observer counted in poor conditions. He saw but 19 percent of the moose present. Effect of Pilot The present experiment attempted to minimize the effect of the pilot on the success of the counter by not allowing the pilot to point out moose he saw. This was necessary to eliminate effects of the pilot's "learning" the moose in a pen after several flights. However, this design differed from actual counting conditions, where the pilot assists in counting game. On the initial flights (1970) in good conditions (Table 3) by each of two pilots, record was kept of total moose seen by pilot and observer, as well as by the observer only (the pilot made mental notes only). These two pairs saw 65 percent and 80 percent of the 49 moose present, significantly (P ( .02; t-test) more than seen by four experienced observers alone the same day. 8 Individual nonobserving pilots (1971) did not significantly (P) .10; t-test) affect proportion of moose seen by observers of any category nor observers' rank within their category (Table 4). Effect of Time of Day Moose surveys are traditionally flown at dawn and shortly after to coincide with mooses' most active period (cf: Geist, 1960; LeResche, 1966; LeResche and Davis, this report Job 1.1). While early morning counts may take advantage of moose activity, lighting conditions may be more conducive to observing moose at other times during the day. Table 5 summarizes mean ranks within counter groups of observers flying before 1000 hours A.S.T., 1000-1200 hours, 1200-1400 hours and after 1400 hours. No statistically significant differences were detected between observers of the same category flying in these time periods. However, the data do suggest that during early spring, moose are more easily seen in mid-morning and after 1400 hours (through about 1530 hours). Effect of Type of Aircraft PA-18-150 Piper "Supercubs" have become the aircraft of choice for aerial surveys in Alaska because of their slow flight characteristics. Canadian federal and provincial (pers. comm.) biologists use other types of aircraft either not generally available in Alaska (eg: Pilatis-Porter; Dornier; DeHavilland Beaver) or too fast for anything but transect work (eg: Cessna 180-185). Some provincial agencies use helicopters in composition surveys of a pre-determined sample size of animals. Helicopters were used with two observers (plus pilot) on three counts of the enclosures under excellent conditions in January 1970 (LeResche, 1970). These observers saw a mean of 75 percent of moose flown over, significantly more (P ( .01; t-test) than seen by single experienced observers in excellent conditions in 1971. However, the cost:benefit rat'io (Bell G4A helicopter:$135/hr; PA-18-150:$35/hr) indicates use of supercubs even when attempting total enumeration of moose. Effect of Terrain and Habitat The four Moose Research Center enclosures are similar in habitat type and therefore it is difficult to ascertain effect of terrain and habitat on proportion of moose seen. Previous experiments (LeResche, 1970) showed significant (P ( .01) differences between proportions of moose seen in all pens with most moose (proportionately) being seen in the pen (Pen 1) with fewest present and fewest being seen in the pen (Pen 4) with most present. Results from 1971 show significant differences only between Pens 1 and 3 (58 percent vs 48 percent; P ( .01; t-test). During experiments this latter year, number of moose present varied less between pens (7-18 in 1970 vs 12-16 in 1971); whereas terrain and habitat remained essentially the same. Thus, number of moose present seems correlated with proportion of animals seen and no difference re: habitat is demonstrable in this rather homogeneous four-square-mile area of seral range interspersed with remnant stands. 9 Table 4. Effect of pilot on ability of observer to see moose, Kenai Moose Research Center, 6-9 March 1971. Inexperienced (A) Experienced/not current (B) Experienced/current (C) Pilot I 43 percent Pilot II 41 percent Mean rank in group by pilot. Pilot I II Number of Observers 18 13 48 percent 70 percent 41 percent 66 percent Counter Group A B c 4.0 3.2 4.8 5.0 4.7 5.0 NB: Pilot-pilot differences not statistically significant (P} .10; t-test). 10 All 3.9 4.9 Table 5. Effect of time of day on proportion of moose seen, Kenai Moose Research Center, 6-9 March 1971. Number of Mean Rank, "Moose Time Counters All Counters Observabili ty" Before 10 a.m. 6 4.7 ± 1.3 23 10 a.m. -12 noon 6 3.2 ± 1.3 31 12 noon -2 p.m. 10 5.7 ± 0.7 18 After 2 p.m. 9 3.3 ± 1.1 30 * 1/mean rank x 100 NB: Mean ranks do not vary significantly statistically (P} .10; t-test). 11 Pellet-count Census Table 6 summarizes pellet groups found in 159 8 x 24' plots in Pen 1 (2-4 June 1971). All groups were deposited between April 1970 and these dates, for plots had been cleared of pellets the past year (LeResche, 1970). The table also includes similar data for 1970, when plots had not been cleared the previous year. In 1970 (LeResche, 1970) an attempt was made in two pens to visibly separate groups deposited during the previous winter. It was estimated that only one-third of pellet groups removed were new and, from this, it was calculated that each moose deposited 10.3 groups per day during a hypothetical 210-day winter period. Current data, from cleared plots in only one-square-mile pen, suggest a defecation rate of 32.2 groups/day. This indicates visual estimates of age of pellets were incorrect and that plots should be cleared to validate the technique (cf: Neff, 1968). Plots will be read in all four pens in 1972, at which time the technique's validity will be substantiated or disproven. Immobilizing Drugs M-99 Etorphine and M 50-50 Diprenorphine: These drugs (immobilizing agent and antagonist, respectively) continued to be most useful for handling animals within the pens. They were the only safe drugs for use on calves, yearlings and rutting bulls. Dosage could be varied to make the animal tractable but still able to walk or immobile (ie: for halter-leading). Multiple doses are safe with no waiting period. Antagonist injected intra- venously usually produced complete reversal of effects within 40-50 seconds. Immobilizing doses varied by season, the animal's age and size. Standard doses ranged from 4-5 mg per 700-800 pound cow in spring to 6-9 mg per 900-980 pound cow in October. Antagonist was administered at twice the dose of M-99. A narcotics license is required to possess the expensive ($1.25/mg) drugs and animals treated with the substances should not be consumed. Succinlycholine chloride ("Anectine"): This drug, in concentrations of 10 mg/cc, continued to be most used because it is easy to secure and cheap, and meat from treated animals is fit for consumption. Therapeutic ratio is very small, however, and some animals (6 of 116) are killed. The drug should be considered unsafe for use on calves, yearlings and rutting bulls. Dosages of 20 mg/adult moose are usually nonlethal but sometimes do not produce immobilization. Multiple doses are almost always lethal unless separated by at least one hour. Do~es of over 23 mg/adult are likely to be lethal. Table 7 summarizes effects of Anectine when administered by dart from helicopter and emphasizes seasonal variability in the drug's effect. The table also demonstrates the effect of hyaluronidase ("Wydase") when mixed with the drug. 12 1-' w Table 6. Pellet groups deposited between April 1970 and June 1971 on 159 8 x 24 foot plots randomly selected by habitat types in Pen 1. Kenai Moose Research Center. Birch S£ruce Mature Hardwoods All Winter Groups Summer Groups Total Groups Plots x Winter x Summer & Winter s 1970 plots (from LeResche, 1970) x Summer & Winter s Dense 32 0 32 25 1.28 1.28 1.37 22 .409 .651 Medium Thin 18 8 2 2 20 10 24 26 0.75 0.31 0.84 0.38 0.94 0.55 25 25 1.00 0.84 1.44 1.08 Spruce-Birch Spruce Dense Thin 8 3 10 14 93 1 0 0 0 5 9 3 10 14 98 23 19 20 22 159 0.35 0.16 0.50 0.64 0.59 0.39 0.16 0.50 0.64 0.62 0.57 0.36 0.82 0.79 0.90 24 20 19 21 157 0.92 0.65 0.21 0.43 0.67 0.95 0.73 0.52 0.58 0.97 196 1-' ~ Table 7. Knock-down times and effects of succinylcholine chloride and 11Wydase 11 administered to free-range adult moose. (Sample sizes in parenthesis). Drug succiny !choline succinylcholine succinylcholine chloride chloride + Wydase chloride 500 ml units Sex M F M F M F Dose/animal: 20 mg 11.7 11.8 (6) (10) 21 mg 7.3 10.4 4 8.4 6.3 7.0 (8) (27) (1) (5) (7) {2) 22 mg 9.1 7.8 6.7 (9) (5) (32) 23 mg 7.6 9.7 (7) (17) % Immobile 65-70 86 65-70 % Killed 4 3 16 Mean knock-down time 7.3 10.1 7.2 7.2 8.0 10.2 ( 8) (36) (6) (37) (20) (29) Both Sexes 9.6 7.2 9.4 (44) (43) (49) In June, when moose reach their lowest physiological state (cf: this report, Job 1.1), immobilization with succinylcholine-chloride is dangerous. The drug was injected into 83 bulls and females without calves in early June. Minimal doses (20 mg) immobilized fewer than 60 percent of moose darted and still killed 11 percent (2 of 18). Dosage sufficient to immobilize 80 percent of moose darted (23 mg) killed 20 percent (8/40). Therefore, the drug was unsuitable for use on animals during this time of year. In contrast, of 72 animals immobilized in March with succinylcholine chloride, only three (4 percent) died--two after rising and moving off. Immobilization proportion when using succinlycholine chloride alone was still low, however. Hyaluronidase ("Wydase"): This enzyme was added to succinlycholine chloride (10 mg/cc) at the rate of 500 nf units per dart (21-22 mg). In May, of 80 animals darted (Table 7), 69 (86 percent) were immobilized and only two of these (3 percent) died from the drug. Knock-down time was decreased 25 percent, from a mean of 9.5 minutes to 7.2 minutes under these free-ranging conditions. Hyaluronidase, by increasing drug absorption, seems to 1) increase the proportion of animals immobilized without increasing mortality and 2) shorten knock-down time. The latter effect is beneficial both in decreasing stress and lowering cost. Under penned conditions, immobilization of trapped animals was hastened even more (ca: 45 percent). Although enough strictly-comparable data are lacking as yet, reduction in knock-down time from ca: 9 minutes to ca: 5 minutes in the same individuals under the·same dosage was common. "Sernalyn" (phencyclidine hydrochloride) plus "Sparine" (promazine hydrochloride) were used on one trapped eight-year-old cow moose with poor results. A total of 1050 mg Sernalyn plus 225 mg Sparine was injected via six darts in a six hour period (600 mg + 150 mg within 45 minutes initially). The animal weighed approximately 850 pounds. Although ataxia and respiratory distress were evident within three minutes of the first 200 mg Sernalyn + 50 mg Sparine dose, the animal was never tractable and had to be roped and tied for dart removal after seven hours. Survival has not been confirmed. "Tranvet" (Propriopromazine hydrochloride): This t.ranquillizer was used on two moose with mixed results. In June a seven-year-old cow, weighing an estimated 800 pounds (360 kg) immobilized with 24 mg succinylcholine chloride, was given 200 mg tranvet. She recovered from the succinlycholine chloride in 32 minutes, but remained approachable to within 10 m (yet able to rise and walk) for 5.5 hours. She was still too alert to handle. She remains healthy. In July a tame yearling bull (weight: 550 pounds -250 kg) was immobilized with 19 mg succinylcholine chloride. Knock-down time was five minutes. At 10 minutes after injection he was given 250 mg tranvet and died at 12 minutes when respiration stopped. Whether death was from 15 succinylcholine chloride only or the combination of drugs is unknown. Since tranvet was given intramuscularly it is unlikely it had significant effects. "Bay-Va 1470" (Bauer-Werke; Belgium): This drug was supplied by Dr. A. Bubenik with a recommended dosage of one mg/kg. It is a CNS depressant. In September a 580 pound (264 kg) yearling cow was given an initial dose of 250 mg. She became slightly ataxic and hyperactive, but could not be roped and tied until she had received an additional 100 mg at 37 minutes and 200 mg at 52 minutes. When down (84 minutes) she had no eye reflex, but good blood pressure and respiration. After four hours in a recumbent position, she rose alertly and ran off, slightly ataxic. She died in the woods 100 m from the trap. Pentabarbital sodium ("Halatal" 1.0 gr Icc): A seven-year-old cow (weight estimated: 850 pounds -390 kg) was given 100 cc of this drug (6500 mg) intravenously after immobilization with succinylcholine chloride, She showed no effects--rising and walking off 37 minutes after immobiliza- tion. She remained healthy therafter. Radio-Telemetry Results of this study are reported in Job 1.2. Freeze-Branding The two attempts were not successful. The 60 second brands created scar tissue and a standard burn-brand, which was covered by hair in summer. Twenty and 40 second brands were not evident after hair regrowth. No unpigmented hair emerged. Other Marking Methods The described pendants, when hung parallel to the mooses' longitudinal axes, were readable from Supercubs more than 90 percent of the time, upon repeated passes. Hung perpendicular to the axis, they could be read fewer than 30 percent of the time. Striped collars were individually identifiable readily from aircraft. Moderate collar loss is suspected, but its extent is unknown as yet. RECOMMENDATIONS Aerial counts of moose should be considered only as trend indicators, never as total counts. To be comparable year-to-year, they should be conducted only under excellent snow conditions, with only currently experienced observers. Hyaluronidase should be used when free-ranging moose are being immobilized with succinylcholine chloride. 16 LITERATURE CITED Bennett, L. J. ~ al. 1940. A study of deer populations by use of pellet- group counts. J. Wildl. Mgmt. 4:398-403. Benson, D. A. 1966. Use of aerial surveys by the Canadian Wildlife Service. CWS occ. pap. no. 3. 35 + 5 p. Bergerud, A. T. 1968. Numbers and densities. In Galley, F. B. & Buechner, H. K. (ed.). A Practical Guide to the Study of the Productivity of Large Herbivores. IBP Handbook No. 7 (Oxford). p. 21-42. Bergerud, H. T., H. Butt, H. L. Russell and H. Whalen. 1964. Immobiliza- tion of Newfoundland caribou and moose with succinlycholine chloride and cap-chur equipment. J. Wildl. Mgmt. 28(1):49-53. Bergerud, A. T. & F. Manuel. 1969. Aerial census of moose in central Newfoundland. J. Wildl. Mgmt. 33:4:910-16. Bishop, R. H. 1969. Moose report. Annual Project Segment Report. Vol. X. W-15-R-3. Work plan K. 152 p. Bowden, D. C., A. E. Anderson & D. E. Medin. 1969. Frequency distributions of mule deer fecal group counts. J. Wildl. Mgmt. 33:4:895-905. DesMeules, P. 1962. Intensive study of an early spring habitat of moose (Alces alces americana) In Laurentides Park Quebec. Paper pre- sented at N.E. Wildlife Conf. Monticello, N.Y. 12 p. mimeo. Evans, C. D., W. A. Troyer ,& C. J. Lens ink. 1966. Aerial census of moose by quadrat sampling units. J. Wildl. Mgmt. 30:767-776. Farrell, R. K., G. A. Laisner & T. S. Russell. 1969. An international freeze-mark animal identification system. JAVMA 154:12:1561-72. Fujita, T. 1970. Delayed hypersensitivity reaction to succinlycholine. Tohoku J. Exp. Med. 102:33-5. Geist, V. 1960. Diurnal activity of moose. Memoranda Societatis pro Fauna et Flora Fennica. 35:95-100. Goddard, John. 1966. The validity of censusing black rhineroceros popu- lations from the air. Reprinted from the East African Wildlife Journal, Vol. 5, August 1967. 7 pp. Harthoorn, A. M. 1965. Application of pharmacological and physiological principles in restraint of wild animals. Wild. Monographs 14:78 p. Harthoorn, A. M. 1968. Manipulation of animals for experimental purposes. In Galley, F. B. & Buechner, H. K. (eds.) A Practical Guide to the Study of the Productivity of Large Herbivores. IBP Handbook No. 7. (Oxford) p. 125-31. Houston, D. B. 1969. Immobilization of the Shiras moose. J. Wildl. Mgmt. 33:534-537. 17 Kambitsch, L. M. Wittman & M. Hemstrom. 1969. Freeze-branding: equipment and methods. U. Idaho Agricultural. Exten. Serv. Current Info. Ser. 100:4 p. LeResche, R. E. 1966. Behavior and calf survival in Alaskan moose. M. S, Thesis, U/Alaska. 84 + xiii p. litho. LeResche, R. E. 1970. Moose report. Annual Project Segment Rept. Fed. Aid in Wildlife Rest. Alaska Dept. of Fish and Game. Miller, F. L. 1968. Immobilization of free-ranging black-tailed deer with succinylcholine chloride. J. Wildl. Mgmt. 32(1):195-97. Neff, D. J, 1968. The pellet-group technique for big game trend, census and distribution: a review. J. Wildl. Mgmt. 32:597-614. Pienaar, U. DeV. 1968a. Capture and immobilizing techniques currently employed in South African national parks and reserves. In Golley, F. B. & Buechner, H. K. (eds.) A Practical Guide to the Study of the Productivity of Large Herbivores. IBP Handbook No. 7. p. 132-44. Pienaar, U. DeV. 1968b. Recent advances in the field immobilization and restraint of wild ungulates in South African National Parks. Acta Zoologica et Pathalogica antverpiensia. 46:17-38. Rasmussen, D. I. & E. R. Dorman. 1943. Census methods and their applica- tion in the management of mule deer. Trans. N. Am. Wildl. Con£. 8:369-379. Rausch, R. A. & A. E. Bratlie. 1965. Assessments of moose calf production and mortality in southcentral Alaska. Ann. Conf. W, Assoc. State Game & Fish Comm. 45:11 p. Rausch, R. A. & R. H. Bishop. 1968. Report on 1966-67 moose studies. Annual,Project Segment Report. Vol. VIII & VIX. W-15-R 2-3, Work PlanK. 263 p. Slater, L. 1963. (ed.) Biotelementry. Pergamon, London. 369 p. Siniff, D. B. and R. 0. Skoog. 1964. random stratified sampling. PREPARED BY: Robert E. LeResche James L. Davis Game Biologists 18 Aerial censusing of caribou using J. Wildl. Mgmt. 28: 391-401. APPROVED BY: State: Cooperators: Project No. : Job No.: JOB PROGRESS REPORT (RESEARCH) Alaska Paul LeRoux; Alaska Department of Fish and Game; U. S. Bureau of Sport Fisheries and Wildlife: Kenai National Moose Range W-17-3 Project Title: Moose Investigations Job Title: Kenai Peninsula Moose Population Identity Study Period Covered: July 1, 1970 through June 30, 1971 SUMMARY Four hundred and thirteen sightings of 283 adult moose tagged on the Kenai Peninsula revealed migratory patterns, concentrating areas, and separate population identities of moose representing an aggregate number of nearly 10,000 animals. Most of the groups studied were seasonally migratory, and moved from lowland wintering areas to calving areas in springtime, thence (in early summer) to upland summering--rutting areas, and back to wintering areas in early-mid winter. This group comprised many large bulls and cows, but very few calves. Sexes were segregated during the early summer migration to the highlands, when males migrated earliest. Sexes inter- mingled during rutting in fall. The other population segment (comprising predominately cows with calves and younger bulls) remained resident in lowland areas of the 1947 burn year-round. Specific drainages appeared to be the sites of rutting by the same individuals year after year. Individuals also followed stereotyped circular migration paths for more than one year. One calving area (the Moose River Flats) was a concentrating spot for individuals from all rutting areas studied (many 60-80 kilometers distant). i SUMMARY •. BACKGROUND OBJECTIVES PROCEDURES FINDINGS . Population Identities Movements •• Concentrating Areas RECOMMENDATIONS. LITERATURE CITED . • CONTENTS BACKGROUND Page No. i 1 2 2 5 5 20 22 22 27 Moose in the lowland areas of the northern Kenai Peninsula receive considerable hunting pressure in the few restricted areas where access exists. In late fall, moose herds in these areas characteristically-have a low proportion of bulls, and trophy-size bulls are extremely rare. Although lowland areas contain a higher proportion of calves within the herd, calf production in some years is lower than anticipated (eg: 33 calves:lOO cows in November, 1970). Most of the area in question is seral birch range remnant from the 1947 burn, and birch browse is in great abundance. However, substantial numbers of moose have died during severe winters in the area. Population estimates by personnel of the Kenai National Moose Range suggest substantial numbers of moose (7900 ± 1400 minimum north of the Kasilof River in early 19 71), but concern has been expressed regarding the numbers and welfare of the "lowland" moose, especially in relation to hunting pressure. The moose traditionally using climax willow ranges in foothills and mountains, but wintering on the lowland areas, receive little hunting pressure. These groups characteristically exhibit a high bull:cow ratio and a low proportion of calves, With the formalization of moose management plans for the Kenai and the designation of certain areas as trophy, foot-hunting and maximum sustained yield hunting areas, delineation of these various groups, their interactions, their seasonal movements, and their calving and breeding sites, has become imperative. Further, the proposed classification of more than one million acres of the area as wilderness, as .well as the possibility of a limited access road bisecting part of the area, requires specific knowledge of the migrations of these moose. Descriptions of populations and their movements would 1) allow harvesting of desired portions of specified moose herds and prevent harvesting of trophy-class bulls while they are away from trophy-management areas (and often antler- less), 2) prevent unnecessary restriction of activities (eg: by wilderness designation) in areas of key winter range, where habitat manipulation might someday become necessary, 3) contraindicate development of small areas 1 seasonally crucial to large numbers of moose (eg: during calving, rutting, or wintering) and 4) provide valid data relative to possible obstructions presented by future proposed highways and other projects. The literature contains few major studies of moose migrations and/or movements, and the studies that have been undertaken have shown that such movements vary with the population studied. Goddard (1970) reported an Ontario study similar to ours. His recoveries were few (59 of 328 marked moose) but he documented movement from summer to winter ranges (done pre- viously by Edwards and Ritcey, 1956; Kraft, 1964 and Houston, 1968) and suggested there was no net movement into heavily hunted areas. Phillips and Berg (1971), with many relocations (2,000) of few (27) radioed Minnesota moose, recorded individual home ranges of 2-10 square miles, winter confinement to less than 100 acres, average daily movement of 0.60 miles, identical mean daily movements of cows and bulls, and 0.5-21 mile movements from winter to summer ranges. VanBallenberghe and Peek (1971) also radio-tracked moose in Minnesota. They showed summer localization, winter confinement by snow, adjacent winter and summer ranges of an individual, and a rapid 12-mile movement by a rutting bull. Mercer and Kitchen (1968) described dispersal of moose introduced onto the Labrador Peninsula. LeResche (1968) and LeResche and Davis (this report; Job 1.2R) reported localization of parturient females and their new calves, and LeResche (1970) suggested internal triggering as a factor in moose migrations. Bishop (1970) reported that a Tanana Flats (Alaska) calf-tagging study suggested that both resident and migratory individuals were present in these lowlands in spring. Didrickson (pers. comm.) reported adult moose tagged in the Matanuska Valley (Alaska) moved nearly 60 miles on occasion. OBJECTIVES To identify populations and key habitat areas and to learn seasonal patterns of movement by moose on the Kenai Peninsula. PROCEDURES Table 1 lists moose marked 1) in October 1968 at Mystery Creek ("highlands"), 2) in March 1970 at Bottenintnin Lake ("lowlands"), 3) in June 1970 at the Moose River Flats ("lowlands"), 4) in March 1971 at the Skilak-Tustumena Benchland ("highlands"), S) in May 1971 at the Moose River Flats, and 6) from August 1969 through May 1971 at the Moose Research Center ("lowlands"). The moose represent, respectively: 1) a rutting group, 2) a wintering concentration, 3) a calving concentration, 4) a late- winter remnant group in a fall rutting concentration area, 5) a calving concentration, and 6) inhabitants of the 1947 burn area during all months of the year. Fig. 1 shows tagging areas and associated geography. 2 Table 1. Moose tagged in GMU lSA, Kenai Peninsula, October 1968-May 1971. Males Females Sex ? Calves Total Mystery-Dike Creek (highlands) 10 18 0 0 28 October 1968 Bottenintnin Lake (lowlands) 16 52 1 0 69 March 1970 Moose River Flats (lowlands) 26 43 2 0 71 June 1970 Moose Research Center (lowlands) 3 40 0 7 50 Moose River Flats 10 51 0 0 61 May 1971 Skilak-Tustumena Bench 2 2 0 0 4 April 1971 Totals 67 206 3 7 283 Male Female Area Collar Ear Collar Ear Pendants Mystery Creek Yellow Left Orange Red Right Orange None Bottenintnin Lake Blue Left Orange White Right Orange None Moose R. Flats ( 19 70) Blue Left Green White Right Green Red Al-AlOO MRC Blue Left Silver White Right Silver White 51-100 Moose R. Flats (1971) Yellow/ Left Yellow Pink/ Right Yellow Red Cl-ClOO orange* red* Skilak-Tustumena Yellow/ Left Yellow Radio Right Yellow Red: "c- Bench land orange* series" * Colored stripes on both sides of collar make the moose identifiable as individuals. 3 ,, . .Moos~ _,,;;;~;-:-~~;,-_,. Pm<it· ! ~~£; r· , f)1 ,', -. IJ;, _ 1 .. ~~~~~. I l'! i~-~ -~"·~~ 1//r ' '· • -~ 1\ll ,.\-:· C!Wifl ;~~./qndj ,;3· .. :'(' . -:,.::::,~,~ /.(/~~; '-~i:-~h-;1~--:, hi! ;/1\NI Ai1 , , r· (;,.('//{~[~'; ' ' -.--~ ~~.~-,L; ." . : I . _:~i.S~epling -· ---~~+··,. .. ('.flU! r· ., ·-·-..:!~,._ Naptowrw ale~ •I ~ ·~"== --~-= __ Jl I I "t: '"''-""'' c•~,-;·n -.·f A,,,. L jM I I I I I . :-t~ ,L,ilcC r 0 ·--., .II• .I ~ • T I 'O- L 'frying Pan·-.. _ &" '''""d <f> -caribou 1: •' ·Islands --· • 4 ;.-· ' ', Moose were tagged using helicopters (or fenceline traps--Group 6) and succinylcholine chloride in projectile syringes. Groups 1, 2 and 6 were ear-tagged and collared to be distinguishable from afar by group and sex but not (except for a few pendant-carrying animals in Group 6) as individuals. Animals in Groups 3, 4 and 5 were made identifiable as individuals by numbered pendants and/or collars and/or color-coded collars. All animals are distinguishable individually when "in hand" by numbered metal ear-tags. Table 2 lists reconnaissance or counting flights made by Alaska Department of Fish and Game personnel. Several additional flights were made by R. Richey of the U. S. Bureau of Sport Fisheries and Wildlife and many resightings of marked moose were reported by the public. FINDINGS Table 3 lists the 413 recoveries and sightings of tagged moose recorded through 15 June 1971. When analyzed by season, location and (tagging) groups, these sightings suggest several facts relative to population identities, movements, and concentrating areas. Population Identities Several biological "populations" (ie: randomly interbreeding groups) are represented by the groups of tagged moose. In one case certainly (Mystery Creek) and at least partially in another (Moose Research Center tagged animals), true populations were tagged as such. In the other sites of concentrated tagging effort, various separate breeding popula- tions apparently were tagged together during nonbreeding aggregations. The Mystery Creek population is perhaps typical of 10-15 separate breeding groups, which gather in separate drainages on the west slopes of the Kenai Mountains north of Skilak Lake (eg: Mystery-Dike, Thurman, Chickaloon, Indian and other creeks) and in the mountain canyons of the Resurrection Creek drainage. This (Mystery Creek) population is the most easily understood of all those groups tagged, both because the group has been tagged longest and because it does represent a true population rather than an aggregation of many. All but one recovery of moose tagged in Mystery Creek in October made between September and October in the two years since tagging has occurred within approximately two miles of the tagging site. All drain- ages of the area described have been searched during this time. This group of moose, then, is very traditional in concentrating in a precise drainage during breeding season. During winter, most sightings of moose from this population have occurred along the Sterling Highway and in the large flatland areas of the 1947 burn to the west of Mystery Creek. During this time of year, Mystery Creek moose intermingle with lowland residents (eg: Moose 5 Table 2. Reconnaissance flights by ADF&G searching for collared moose. Date 26 March 19 70 31 March 1970 3 April 1970 6 April 1970 6 April 1970 6 April 1970 8 April 1970 14 April 1970 14 April 19 70 22 April 19 70 24 Apri 1 19 70 27 April 1970 4 May 1970 4 May 1970 11 May 1970 11 May 1970 17 May 19 70 1 June 1970 1 June 1970 23 June 1970 23 June 1970 10 July 1970 17 July 1970 29 July 1970 4 August 1970 10 August 19 70 22 August 19 70 24 August 1970 1 September 1970 1 September 1970 5 October 19 70 5 October 1970 22 October 1970 22 October 19 70 18 November 1970 19 November 1970 23 November 1970 30 November 1970 30 November 19 70 1 December 1970 2 December 19 70 2 December 19 70 3 December 1970 Area Skilak Lake N. of Kenai R. Same Same Same South of Kenai River to benchland North of Sterling Hwy. 0-5 miles Skilak Lake area Moose R. Flats Skilak Lk. area Same Moose R. flats, Hidden Lk. Skilak Lk. Moose R. flats to Sterling Hwy. Mystery Creek Skilak Lk. area Moose R. flats; upper Funny R. Skilak Lk. area Tustumena-Skilak benchland + Skilak area Moose River flats Skilak Lk. area Moose R. flats Mystery Creek Moose R. flats, benchland Mystery Creek Mountains N. of Mystery Creek Moose R. flats, Skilak Lake area Bench land Mystery Creek Moose R. flats MRC Resurrection Creek Bench land Mystery Creek Skilak Lk. Bench1and MRC Juneau Creek Resurrection Creek Skilak Lake Mystery Creek 1947 Burn N. of MRC Thurman Creek, Fuller Lk., Mystery Creek Skilak Lake Skilak Lake 6 Collared Moose Located* 12 BL 10 BL 6 BL 4 BL 0 3 BL, 1 MRC 4 BL 1 MRC 7 BL 6 BL 0 0 0 6 BL 0 6 BL 0 1 MC 1 BL 3 MRF 1 MC 2 BL, 2 MRF 7 MC, 3 MRF 7 MRF 2 MRF 1 BL, 1 MRF 1 MC, 1 MRF 1 MRF 1 MRF 3 MRF 2 BL 4 MC, 3 MRF, 1 BL 3 BL 5 BL 1 MRC 4 MRF, 1 MC 3 MRF 1 BL 1 MC, 1 MRF 3 MRC, 1 MC, 1 MRF 5 MRF, 2 MC, 1 BL 3 BL, 1 MRC 4 BL, 1 MRF Table 2. (cont'd.) Reconnaissance flights by ADF&G searching for collared moose. Date 3 December 1970 4 December 1970 21 December 1970 21 January 1971 21 January 19 71 21 January 19 71 12 February 1971 23 February 19 71 30 March 1971 24 May 1971 25 May 1971 27 May 1971 27 May 1971 15 June 1971 * Code: Tagged at Tagged at Tagged at Tagged at Area Bench land West of Skilak Lake Skilak Lake Skilak Lake Moose R. flats MRC MRC, Moose R. flats, Skilak Lk., Benchland Moose R. flats, MRC area Benchland, Skilak Lake, Mystery Creek Moose R. flats Moose R. flats Moose R. flats Moose R. flats Moose R. flats Mystery Creek: MC Bottenintnin Lake: BL Moose River Flats: MRF Moose Research Center: MRC 7 Collared Moose Located* 1 BL 1 BL 6 BL 1 MRF, 2 BL 1 MRF 1 MRF 0 3 MRF, 1 MRC 1 MRF 2 MRC, 1 MRF 2 MRF 2 MRF 3 MRF, 1 MRC 25 MRF, 1 MRC Table 3. Recoveries and sightings of collared moose, Kenai Peninsula, through June 1971. MONTH Tagging Site I-II III-IV V-VI VII-VIII IX-X XI-XII Totals Mystery Creek 21 FF 14 14 12 7 16 87 1 MM 1 14 6 5 1 115 22 15 31 18 12 17 115 Bottenintnin Lake 4 FF 51 15 19 20 19 128 2 MM 9 2 0 1 1 15 6 60 17 19 21 20 143 Moose Research Center 3 FF 4 7 1 6 5 26 OMM 1 4 1 4 0 10 3 5 11 2 10 5 36 Moose River Flats 6 FF 4 49 11 7 8 85 4MM 1 8 8 2 11 34 10 5 57 19 9 19 119 Table 4. Proportion of sightings outside of four contiguous townships from tagging site, Kenai Peninsula, 1968-15 March 1971. Tagging Site FF % MM % N Mystery Creek 35 43* 14 50* 109 Bottenintnin Lake 34 27 2 13 141 Moose Research Center 7 30 2 22 32 Moose River Flats 16 29 21 72* 84 * Tagged in concentrating areas 8 Tagging Site Mystery Creek Moose River Flats Skilak Loop (Bottenintnin Lake) 1947 Burn (Moose Research Center) Table 5. Seasonal locations of tagged moose 11 populations," Kenai Peninsula. Winter Sterling Hwy. East of 1947 Burn and 19 4 7 Burn West 1947 Burn West, Moose River Flats and Sterling Hwy. West ? 1947 Burn Late Winter Sterling Hwy. East of 1947 Burn and 1947 Burn West 1947 Burn West and probably Skilak Loop Skilak Loop- Naptowne 1947 Burn Calving Moose River Flats and Kenai River Moose River Flats Skilak Loop- Naptowne and probably Brown's Lake Area 1947 Burn and Moose River Flats Summer Sterling Hwy. East and Mystery-Dike Basin Kenai Mts. North, Bench- land and Moose River Flats Benchland and Below 1947 Burn Rut Mystery-Dike Basin Kenai Mts. North and Bench land Benchland and Skilak Loop and Kenai Mts. North 1947 Burn, Moose River Flats-Skilak Loop Post-Rut Mystery-Dike Basin and Sterling Hwy. East and West Kenai Mts. North and 19 4 7 Burn West Skilak Loop- Naptowne and Bench land 1947 Burn Research Center tagged animals) and moose from other Kenai Mountain drain- age breeding groups (Table 6). Mystery Creek moose typically calve along the Kenai River. Bulls inhabit the Moose River Flats during calving. Table 4 and Fig. 2 illustrate the circumstantial data for the identity of this population, and Table 5 summarizes its locations during 6 two-month periods. As shown in Table 4, most observations throughout most of the year are far from the tagging (breeding) site. All sightings during September-October were near the tagging site. Thus, this population concentrates on the breeding grounds. Fig. 2, which represents the mean distance of recoveries from tagging (October) site of Mystery Creek moose by month, shows differential male and female migrations to and from the breeding grounds. Aggregation of both sexes during Septerrnber-October--ie: demonstration of population status--is evident. Tables 4 and 5 and Figs. 3, 4 and 5 present similar data from sight- ings of moose tagged at Bottenintnin Lake (Skilak Loop), the Moose River Flats and the Moose Research Center. Table 4 shows that the Moose River Flats was a concentrating area for bulls from other areas when tagging occurred there (June) , for 72 percent of subsequent sightings of these bulls were more than six miles from the tagging site. Figs. 3, 4 and 5, like Fig. 2, imply certain things about the popu- lation status of the groups in question. Since these groups were tagged outside of their breeding areas, breeding area was "estimated" by circum- scribing the smallest polygon about all September and October sightings of moose from each tagged group and taking the polygon center as the breeding ground. Mean distance from this center, during September-October, thus becomes a measure of group dispersal during the breeding season. A widely dispersed group obviously represents several breeding populations. In addition, difference in this distance between males and females further illustrates intersexual mixing of animals from within the tagged group during rut. For example, Fig. 4 shows that both males and females (tagged at the Moose River Flats in June) are dispersed an average of more than 10 miles from the "center" of their breeding range. In other words, they are not concentrated on one breeding ground and the group concentrated at Moose River in June represents more than one breeding population. Since males and females in this group had approximately equal dis- persals from the breeding center, we may infer that at least some males and females from the group do breed with one another (that moose were tagged from some true populations) . Sightings plotted on a map confirm this. Fig. 3 shows that the group tagged while wintering in the Skilak Loop area likely represents individuals from many populations. In contrast, Fig. 5 implies that males and females tagged at the Moose Research Center are from a true breeding population concentrated near the tagging site (see Table 4). Figs. 6-9 present seasonal sighting locations of moose tagged at Mystery Creek, Skilak Loop (Bottenintnin Lake), Moose River Flats and the Moose Research Center. These figures were used to construct Table 4. 10 Area Bench land Kenai River upstream of Skilak Lake Mystery-Dike Creek Basin Kenai Mts. north of Mystery Creek Moose River Flats 1947 Burn north and west Skilak Loop- Naptowne Table 6. Groups of moose seasonally occupying various Kenai Peninsula areas. Winter Mostly Mystery Creek breeders Late Winter Mostly Mystery Creek breeders Calving Mostly Mystery Creek breeders A few Mystery Creek breeders Sunrrner Bot. Lake winterers, MR Flats calvers Mostly Mystery Creek breeders Rut Bot. Lake winterers, MR Flats calvers Mystery Creek, Bot. Lake, '47 Burn Mystery Creek breeders and Moose River Flat calvers Post-Rut A few Bot. Lake winterers Mostly Mystery Creek breeders Mystery Creek breeders, MR Flat spring, Bot. Lake winterers Moose River Flats calving population Residents Moose River, Mystery Creek and '47 Burn calving grounds Residents Residents Moose River Flats calvers Residents Bot. Lake winterers Benchland breeders and some Moose River calvers Benchland breeders Residents Residents Moose River Flats calvers Bench land breeders and some Moose River calvers <fJ J..u _J - Figure 2. Mean distance from tagging site by month of 109 sightings of moose tagged in Mystery-Dike Creek Basin in October 1968. ~ 10 I Lu l!l ~ D 141----· ··-----· MALES FEMALES ~c2.B 8( 1-2. ~-4 .5-G 7-a 9-1{) ll .. f2 ---------MQN1l4 --------- 12 C/J tu _J -~ ~ ~ ' ~ ~ ~ ~ \ll 0 Figure 3. Mean distance from the center of a polygon enclosing all September -October sightin s by month f 141 sight ngs of moos tagged in kilak Loop rea in &.rch 1 70. ~15 tl' I 30 I I I 126~ 25 I\ I I \ I I I 20 \ I \ I y / 15 / 10 5 1-2 ~-4 5-6 7-8 9-ID 11-12 _________ MOI\.1TH -------- 13 Figure 4. Mean distance from the center of a polygon enclosing all September -October sightings by month of 84 sightings of moose tag ed on the M ose River F ts n June 197 • 2.0 15 10 5 /' I \ -...../ ' -2 ~ -ID II -12 !l-6 '7-S ________ MOI\lTI-1 ------- 14 I I I I I I I I I I I I I I I I I I I Figure 5. Mean distance from the center of a polygon enclosing all September -October sightings by month of 32 sightings of moose tagged at the Moose Research Center (1 47 Burn) ye r-round in 969 -1970. /C 1-2 I I I I I I 1\ I \ I \ I \ I \ I \ I \ I ~-B ?-B \ \ \ \ \ .,____-+-q ~ \ \ 9-ID II --12 --------------------------------------~NT~---------------- 15 Figure 6. I~ \0 Monthly locations of moose tagged in Mystery-Dike Creek Basin in October 1968. OBSERVED AT: D Mystery -Dike Creek Bas in Kenai River East of Ski 1 ak Lake Moose River Flats .......... . . . . . . . . . . •. ·····················: ······················ .......... ····················· ........... ~-1& 7-8 ________ MONTI-\ 16 Skilak Loop - Naotowne Kenai Mountains North of Mystery Creek Basin 9-ID II -12 Figure 7. Monthly locations of moose tagged in Skilak Loop area in March 1970, OBSERVED AT: I ) Skilak Looo -NantaNne l[[[[[[[[[[~[[[[[[[[[[[[[[[[[[[[jjj\JI 19 4 7 Burn I·~~ Kenai Mountains North of I II Mys terv Creek Basin 10 .:5-4 7-8 I I -12 _________ M~~TH _______ _ 17 (/J 4.!i z ,_ :z: \!1 -til u.. (] ~ C(l ~ :l 2 Figure 8. Monthly 'locations of moose tagged on the Moose River Flats in June 1970. 25 20 15' 10 OBSERVED AT: [ I Mystery-0~ ke Creek ,._ ____ _,_ Bas1n r~oose River Flats 1947 Burn (MRC) 1111111111111111111111111!1111111~ Ski 1 ak ILooo - Naotowne Kenai Mountains North of r~ystery Creek Basin ~~~~~(~(~~~~ Skilak-Tustumena Bench 7-8 ~-10 ---------MOI\JTH _________ _ 18 Figure 9. Monthly locations of moose tagged at the Moose Research Center (1947 Burn) year-round 1969-1970. OBSERVED AT: ~Moose River Flats 1947 Burn 10 5 __________ I'V\0"-JTI-4 --------- 19 Primarily, they illustrate group dispersal and reaggregation at the tagging site during the season of tagging. For example, Fig. 8 shows that moose aggregated at the Moose River Flats calving grounds in May- June (5-6) disperse to many areas during other months. In the July- October (7, 8, 9, 10) period moose tagged in this group were seen 1) on the Moose River Flats, 2) in various Kenai Mountain drainages, 3) on the Skilak-Tustumena benchland, and (a special case of (1) above) in the Mystery-Dike Creek basin. The other figures illustrate similar aggrega- tions and dispersals by the other tagged groups. Areas represented on the figures are large (for example, "Kenai Mountains north of Mystery Creek Basin" includes more than 20 drainages and more than 450 square miles--2180 square kilometers). Therefore, aggregations apparent on the figures may be exaggerated. The areas do represent the major physio- graphic areas of the northern Peninsula, however. Further, they adequately define areas suitable for separate management practices. Movements Much movement information is implicit in the above discussion of population identity and the following account of concentrating areas. Seasonal migrations, their year-to-year differences and individual move- ments merit special mention, however. A major migration, as suggested above, occurs from the Moose River Flats (and other scattered calving areas) to the Kenai Mountains and the Skilak-Tustumena benchland. This movement typically occurs from July through September-October. Fig. 10 defines the movement by plotting mean altitude of males and females by season. Sexes are partially segregated from May through August because males migrate to high country earlier than do females, which remain on calving lowlands longer. Further, males migrate to the very heads of mountain drainages, but females typically do not move so high. Intermixing occurs at breeding (months 9-10). After rut, males typically return to upper drainages as females begin migrating to lowland wintering areas. Most males join them in January-February and the sexes remain mixed through June, when the upland migration once again begins. Some males remain near timber- line all winter. There are important exceptions to the above pattern. A substantial portion of the population tagged at the Moose Research Center (in the 1947 burn adjacent to the Moose River Flats) is resident in that area. This is suggested by the sightings of Moose Research Center tagged individuals (cf: Table 4, Figs. 5 and 9) of which 33/36 (92 percent) have been within four miles of the tagging site. It is further suggested 1) by trapping success of traps outside the Moose Research Center fence- line, which is uniformly high during all seasons, 2) by the average of more than 15 moose per square mile enclosed by building the Center's four enclosures (two completed in January, two in August) and 3) by observations adjacent to the Moose Research Center during all months of the year. 20 ~ w lL 1 w c ~ -~ <( ~ ~ Figure 10. Mean altitude by month of 348 sightings of collared moose; Kenai Peninsula. 1968. March 1971. • • FEf\AALE~ 74 Observa ions --TOTAL: 348 Observatio MALES 76 Observa ions 2500 \ ;\ 2000 I \ I \ I \ I \ I I \ I \ 15'0Q I I I 1000 I ' / 5t:£J 1-2 3-4 5·6 7-8 I 1-12 -----------------~NtH ________________ __ 21 Concentrating Areas Concentrations of moose occur during calving, rutting and late winter. On the northern Kenai Peninsula, greatest numbers are most concentrated during calving and rutting. Wintering areas are so vast (ie: the 1947 burn of more than 350,000 acres) that winter concentrations, though impressive (ca: 500-1000 animals in a township near Skilak Lake in March 1970), do not occur to such an extent as calving and rutting con- centrations. Table 6 summarizes present data re: which moose occupy which areas of the northern Kenai during which seasons. This table is organized by tagging group rather than by breeding population because of the limitations of group tagging. Fig. 11 graphically presents the data summarized in Table 6. Tallest bars represent areas where moose are most concentrated during each two-month period. They show concentrations 1) on the Skilak Loop area during March-April (largely of animals tagged there--resighted shortly after tagging) , 2) on the Moose River Flats during May-June (moose tagged there and at Mystery Creek) and 3) on the Skilak-Tustumena benchland and Mystery-Dike Creek basin and other Kenai Mountain drainages from July through October or December (moose tagged in Moose River Flats and Mystery Creek). RECOMMENDATIONS Reconnaissance flights throughout Subunits 15A and 15B should be continued weekly to derive maximum information from tagged animals. More moose should be tagged in late winter concentration areas near the Sterling Highway to adequately define calving and rutting areas of these animals. Moose should be tagged in fall in the Skilak-Tustumena benchland to determine wintering areas of this trophy-class population. Moose should be tagged in fall in one Kenai Mountain drainage in addition to Mystery Creek to determine if the Mystery Creek population is typical. In the interim, seasons should be set with the following probabilities in mind: 1. Late summer and fall "highland" populations are likely older animals (LeResche, 1970) and fewer in number than the substantial lowland resident populations. 2. Populations are predictable in their movements; therefore, harvest of various groups can be controlled by properly timed field announcement hunts. Tables 5 and 6 can serve as guidelines for such harvests. 22 Figure 11. Moose present in seven major areas of the Kenai Peninsula by month, as surmised from sightings of tagged animals, 1968 -1970. 12 6 TAG4en Ar: ___ ]Skilak Loop -~,~~ Moose River Flats 1////////////J//(/////////////////////]J Mystery Creek -1947 Burn (MRC) !18SEA.Utll A-r St1U.Il - lii!:.TliiiFN A MNCJI LAIJ f1 t:JS~f/IJEIJ Ar lleAJAI 111oel EA5T oF ~ILAK LAKE \-2. :?l-4 5-6 1-8 9-IQ 11-12. ______ MONTI-I _______ _ 23 Figure 11. (cont'd.) Moose present in seven major areas of the Kenai Peninsula surmised from sightings of tagged animals, 1968 -1970. 54 :ao .24 18 12. ~tr&En. AT; ooou<.'""" '-"''""' " I <: k; 1 ak Loon Moose Ri ve r Fl ats Mystery Creek 1111111111111111111111111111111111111111 19 4 7 Burn ( MRC) !JBSeii.VElJ 4r StiLAi LtJtiP '-2. 3-4 5-6 1-8 q· .. t 0 II ·12.. ______ MONT~------~-- 24 ~ - Figure 11 (cont 1 d). Moose present in seven major areas of the Kenai Peninsula surmised from sightings of tagged animals, 1968 -1970. \2 6 12 771CrGtn A I: I lllllllllllllllllllllllllllllllllllll\11 Mv~~ll.v -D111~ 7311$/lf/ Ski 1 ak Loop Moose River Flats · Mystery Creek l185ElVEll AT ~AJAI liN. MVSTEA.V CAUN. 'BMIAJ 3-4 25 \I -12 Figure 11. (cont'd.) Moose present in seven major areas of the Kenai Peninsula as surmised from sightings of tagged animals, 1968 -1970. 42 36 !0 IS 12 ll8SER.VEn AT Moos.e l.vEl fLAT~ 5-G 1-8 TAbf:,G1J AT! I lllllllllllllllllllllllllllllllllllllll Skilak Looo Moose River Flats M.vs te ry Creek 1947 Burn (~1RC) -------MONTH _________ _ 26 3. Even though migratory populations may be adequately harvested every several years (on a maximum sustained yield basis) local resident populations in the 1947 burn remain essentially unharvested with present access. 4. Fall rutting populations in mountain drainages are probably composed of the same individuals year after year. Therefore concentrated hunting in certain drainages should be avoided. 5. Migration routes of many (perhaps 3,000-5,000) moose cross the Moose River lowlands in an east-west direction. A barrier (eg: a fenced highway) across this route could be disastrous to these animals. 6. Moose breeding in the Skilak-Tustumena "bench land" (trophy- hunting area of Game Management Unit 15B) utilize the Skilak Loop- Sterling Highway area for wintering and the Moose River Flats for calving. These areas should be managed to preserve maximum numbers of trophy moose during these times of year. LITERATURE CITED Bishop, R. H. 1970. Changes in composition of the Tanana Valley moose herd. Paper presented 6th N. Am. Moose Conf., Kamloops, B. C. Edwards, R. Y. & R. W. Ritcey. 1956. The migrations of a moose herd. J. Mammal. 37:486-494. Goddard, J. 1970. Movements of moose in a heavily hunted area of Ontario. J. Wildl. Mgmt. 34:439-445. Houston, D. B. 1968. The Shiras moose in Jackson Hole, Wyoming. Grand Teton Natural History Assoc. Tech. Bulletin No. 1. 110 p. Kraft, A. 1964. Management of moose in a Norwegian forest. Norwegian State Game Research Inst. Series 2: No. 16. 61 p. LeResche, R. E. 1968. Spring-fall calf mortality in an Alaska moose population. J. Wildl. Mgmt. 32:953-956. LeResche, R. E. 1970. Moose Report. Fed. Aid Wildl. Restoration Proj. Seg. Report. Vol. XI. W-17-2. Mercer, W. E. & D. A. Kitchen. 1968. A preliminary report on the exten- sion of moose range in the Labrador Peninsula, 5th N. Am. Moose Conf. Kenai. 24 + 8 pp. (mimeo). 27 Phillips, R. L. & W. E. Berg. 1971. Home range and habitat use patterns of moose in northwestern Minnesota. 7th. N. Am. Moose Conf. (abstract). VanBallenberghe, J. & J. M. Peek. 1971. Radio telemetry studies of moose in northeastern Minnesota. J. Wildl. Mgmt. 35:1:63 PREPARED BY: Robert E. LeResche James L. Davis Game Biologists APPROVED BY: Research Chief, Division o 28 Game