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TECHNIQUES OF HUSBANDRY IN NORTH AMERICA Charles C. Schwartz Alaska Department ofFish and Game, Moose Research Center, 34828 Kalifomsky Beach Road, Soldotna, Alaska 99669 ABS1RACI': Moose (Alces alces) are maintained in captive or semi-captive conditions in North America for display and education, scientific research, and commercial breeding. Moose husbandry techniques are widely scattered and in most cases poorly documented. In this review, I summarize husbandry techniques obtained from a survey ofmost facilities in North America with moose. In addition I summarize our experiences from the Moose Research Center where a large number (15 - 32) of moose are kept. Detailed descriptions of physical facilities, techniques for feeding and care ofadults and calves, and maintenance of herd health are presented and discussed. Minimum facilities require at least a 2.13 m woven wire fence and shelter. With adequate shelter, moose can toletate extreme cold and wind, but warm tempetatures impose stress; shade and cooling ponds or sprinklers are important. A formulated ration meeting nutritional and physical requirement of moose has simplified feeding and reduced labor costs. Most moose are supplemented with fresh cut browse and other green _plant material. Disease, particularly in calves, can result in high mortality. Moose do not survive in captivity as long as they do in the wild. ALCES SUPPLEMENT 1 (1992) pp. 177-192

Speidel (1965:88) in his review on the METHODS care and nutrition of moose stated that, "Para­ I used 3 methods to locate facilities in doxically this largest and most powerful North America that kept moose in captivity. member of the deer family is difficult to keep First, I contacted all the major listed in in captivity and this poses a challenge to any theNationalZooDirectoryforNorthAmerica. ." Such was the case when we began Second, I requested a list of facilities keeping worldng with captive moose at the Moose Alces alces listed with the International Spe­ Research Center (MRC) in 1977. Research cies Infonnation System. And third, I con­ being conducted at the MRC required a herd tacted moose specialists in all jurisdictions of of tractable moose maintained under captive the United States and Canada where moose conditions. A review of the literature and occur. I requested the names and addresses of correspondence with zoos and other research zoos, wildlife parks, game ranches, and re­ facilities revealed that keeping moose under search facilities that kept moose. Each facility captive conditions was difficult (Schwartz et contacted was asked if they kept moose and, al. 1980). A major obstacle at the time was a if so, to cooperate in a husbandry sutvey. lack of a suitable ration. Unless moose were fed browse, they inevitably died of chronic RESULTS wasting resulting from dianhea. Within the past decade, our understand­ I contacted 121 different zoos, research ing of the nutritional and environmental re­ facilities, and game fanns, of which 100 re­ quirements of moose has flourished sponded. With the exception of a few game (Franzmann et al. 1987, Hjeljord 1987, fanns in , most of the 21 non-respond­ Schwartz et al. 1987a). A small number of ents were zoos located in parts of North moose are now maintained throughout North America well outside the geographic range of America under a variety of captive and semi­ moose. I did not send a follow-up letter. Of captive conditions. The intent of this paper is the 100 facilities that did respond, 29 kept to review current husbandry techniques used moose and all agreed to participate in the in North America. sutvey. I mailed a second questionnaire to all 177

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29 facilities with moose. Twenty-six com­ and bulls) down to $1,200. One person pro­ pleted the questionnaire, 2 did not respond, viding separate prices for male and female and at 1 facility the owner did not speak calves. The average sale price listed for a calf English and was unable to participate. I sent was $500-1000 less than that for a cow. follow-up letters to the two non-respondents, but Bear Country USA, and Wildlife Safari Physical Facilities chose not to participate. Fence~There was much diversity in the There was considerable variation in the size and make-up of the physical facilities detail with which each respondent answered used to keep moose. Fencing used was either each question. For this reason, sample sizes chain link (35%) or woven wire (65%). The varied among questions and responses. Also, most common type of woven wire was 5 X 10 there was variation in experience and the em or 15 X 15 em squares in 12 or 9 gauge duration that the 26 facilities kept moose. The wire. No facility used electric fence. longest history spanned 52 years, and the Fence height ranged from 1.8-3.1 m but shortest only 1 year. Nine facilities kept 2.1-2.4 m fences (92%) were by far the most moose >20 years, 10 10-20 years, and 7 <10 common. The facilities with a 1.8 m fence years. indicated they had moose jump out. Three The primary reasons for keeping moose facilities located in areas with naturally oc­ were display and education (n =14), wildlife curring wild moose and deer( Odocoileus spp) research (n = 9), and breeding on game fanns indicated that they occasionally have wild (n = 3). In addition to display, 3 facilities moose or deer jump into their pens. Two had raised orphaned calves as a service to their 2.1 m fences, and 1 had a 2.4 m fence. One local Wildlife Department. No respondent facility eliminated moose jumping in by indicated they raised moose for meat pro­ putting a top rail around their holding pen. duction. Post spacing ranged from 2.4 m to 7.3 m, Thirteen facilities listed A. a. andersoni, 10 but most were 3-5m apart. Galvanized metal A.a.americana,SA.a.gigas,and 1A.a.shirasi or 10 X 10 em treated wood posts was the most as the subspecies ofmoose kept; 4listed more common type listed. Only 5 (20%) facilities than one subspecies. All but one facility used a top rail. A few facilities indicated that obtained their original stock from the wild, they had guard rails parallel to the ground at usually within the state or province where the varying heights (0.6-1.8 m) above the ground. facility was located. Only 2 facilities (6%) Most respondents liked their fences. One have kept moose for 5 generations; most have facility indicated that on 2 occasions they had kept moose for only 1 (35%) or 2 (30%) young calves get tangled in the 15 X 15 em generations. Most facilities routinely sup­ woven wire mesh when they became fright­ plemented their captive stock with animals ened and ran into the wire. A few individuals from the wild. indicated the lighter gauges of wire were Nine facilities provided sale prices of easily damaged by the antlers of bulls. We moose and 2listed purchase prices. Prices for have had similar experience at the MRC. One bulls averaged $1,500 (all prices are in 1990 facility indicated that they have had animals USdollars),andrangedfrom$1,000to$5,000. chew on wooden posts and other furnishings Females older than calves averaged $3,500, within the pen. They felt that this was a result with the low price listed at $1,500: the top of a lack of adequate browse. Two facilities price listed was $6,500. Calf prices were the had occasional problems with black bears most variable, ranging from $7,000 (this (Ursusamericanus) and (Canis lupus) person did not provide sale values for cows digging under their fences.

178 ALCES SUPPLEMENT 1 (1992) SCHWAR1Z- MOOSE HUSBANDRY

There were several suggestions given to weather, they stress easliy in heat and humid­ improve fencing design. Probably the most ity. During the summer, moose begin to show common was to put the fence wire on the signs ofheat stress when temperatures exceed inside of the posts. This provided a smooth 14-20 C (Renecker and Hudson 1986). fence line within the pen, and prevented bulls Cooling facilities (n = 24) included shelters from "popping fencing staples" when they (38% ), ponds or streams (50%), trees (50%), pushed on the wire. One facility attached an and sprinklers and misters ( 17% ). Most (71%) apron of fencing 91 em wide which was had more than one type of cooling facility. buried. This apron effectively kept wild Facilities located south ofnonnalmoose range predators and dogs from digging into the pen. used sprinklers. Most people indicated those One respondent felt sliding gates facilitated moose seek cool places, particularly water, easy animal handling. whentemperaturesreach27Cand/orhumidity At the MRC, we tested the New Zealand is high. No facility indicated heat stress to be type electric fencing (Gallagher-Snell Power a major problem. Fence, San Antonio, Texas) which is high Several people provided valuable sug­ voltage (7 ,900 V). The fencing worked and gestions to improve holding facilities. In successfully confined moose, but required smaller enclosures, protection of trees from much maintenance, particularly when we kept overbrowsing and physical damage was im­ antlered bulls in the pens. From our obser­ portant. Bulls need a few unprotected trees vations, it appeared that hardened antlers did and shrubs to rub velvet from antlers. not always conduct electricity. We watched A hard concrete surface near the feeding bulls touch the electric wires with their antlers area was useful to prevent muddy conditions and show no indication of a shock. As a from developing in this high use area. Con­ consequence, one male pulled down part of crete also provided a rough hard surface that the fence wire. Animals touching the fence ensured adequate hoof wear. Several people with long winter guard hairs occasionally who kept moose on soft substrates indicated received no shock. Snow acted as an insula­ hoof growth problems. Conversely, one fa­ tor, therefore positive and negative wires were cility indicated excessive hoof wear due to a alternated on the fence to assure adequate concrete floor in their shelter; the moose liked grounding of animals touching the fence. this shelter and spent a considerable amount Shelter-Moose are well adapted to the of time in it northern environment and with adequate pro­ Stockingrates-Stockingrates varied among tection from wind they can tolerate very cold facilities, with as few as 1 moose/60 ha when temperatures ( -30 C) (Renecker and Hudson moose were consuming natural forages to as 1986). Fourteen (64%) of the facilities pro­ many as 1 moose/0.1 ha when they were on vided theirmoose with a winter shelter. Barns, display and fed a fonnulated diet When block houses, or 3-sided shelters were the moose were stocked at a high density, the most common. No facility used a heated facilities in all cases kept fewer than 3 moose shelter. Eight (36%) provided no shelter but but even then most facilities maintained a the holding pen contained dense stands of density of about 1 moose/0.5 ha. At the MRC, coniferous trees. No respondent indicated we have maintained a mixture ofcows, calves, winter weather caused their animals prob­ and bulls in groups ranging from 5-15 indi­ lems. One facility used their shelter as a viduals in a 4 ha pen during fall and winter. holding and switching area. The pen contains several areas ofdense spruce Although moose are capable of maintain­ cover allowing animals to segregate. With ing body temperatures in extremely cold this number of animals, we did witness ag-

179 MOOSE HUSBANDRY- SCHWARTZ ALCES SUPPLEMENT 1 (1992) gression, especially in .the area where the (Franzmann et al. 1975, Renecker 1987). animals were fed the pelleted ration. The volume of replacer fed at each feed­ All but 2 facilities kept bulls mixed with ing is important Excessive feeding can result cows. The two not keeping males reared only in diarrhea, listlessness, potbellies, and death. females and one of these facilities imported a Neonatesarerarelycapableofregulatingmilk bull for breeding. One facility kept bulls intake (Robbins 1983). Addison et al. (1983) separated from females from calving to felt that excessive diarrhea in moose calves breeding season. Four research facilities re­ raised in was directly related to the moved the antlers from males: no facility volume of replacer fed. They recom­ displaying moose removed antlers. One zoo mended that calves not be fed more than 2.5 V also removed antlers from animals that were day, and volume increases in fonnula should shipped. Rutting males can cause problems not exceed 10-15%/day. Similarly, avoid including destruction of fences, feeders, and underfeeding. trees. Several people cautioned about the Often the best indicator offeeding level is dangerous nature of male moose during the the health and vigor of the calf. Rates of rutting season. Only one facility indicated weightgaininhealthycalvesarearound 1.5%/ that an antlered male had injured a female. A day(Schwartzetal.1987b)whichisabout0.3 3-year-old bull gored a yearling female at the kg/day in young calves and as much as 1.4 kg/ MRC. This female died from the injuries. day in late summer calves. Eighty-six percent of the facilities bottle­ Feeding and Care raised calves. Eighteen facilities raising moose Calf Raising-Several references in the calves, including the MRC, provided usable literature address bottle-raising of neonatal data summarizing their techniques (Table 1). moose calves (Addison and Dodds 1959, Diets varied but most (44%) facilities used a Yazan and Knorre 1964, Markgren 1966, mixture of whole cow's milk and evaporated Regelin et al. 1979, Addison et al. 1983, and milk. Mixtures ranged from 5 to 1 part(s) Welchetal.1985). Eachdiscussesadifferent whole (4% fat) homogenized cow milk per technique. part of (Table 1). Six (33%) Robbins (1983) provides a good basic list of the facilities used commercial cow milk of items to consider when preparing a for­ replacers (various brands) either alone or mulated milk replacer for raising orphans. mixed with whole cow or milk. Three Milk fonnulation is important. Proper bal­ (16. 7%) used evaporated milk diluted with ancing of sugars and fats of similar compo­ water, and one facility used a product called sition to maternal milk is desirable. If fat is doe milk replacer (National Food Laboratory, added, is preferred but animal tallow Inc., Dublin, CA 94568). Doe milk replacer is oreggyolkscan also be used. When significant similar to deer ( Odocoileus spp) milk, and levels of fat are added, homogenize the milk contains about 29% fat compared to cow milk to ensure maximum fat dispersion and mini­ replacer that has about 13.5% fat. Reconsti­ mal fat droplet size. These measures will tuted doe milk contains approximately 6.8% maximize digestibility. Added fats that pro­ crude fat, 6.8% crude protein, and 23% solids. duce severe diarrllea include vegetable fats, Several people added egg yolks, ­ primarily common cooking oils. Avoid milk, casein, and/or a commercially available feeding milk higher in fat content than the product called Multi-Milk.TM (Pet-Ag. Inc., species nonnally produces. Moose milk con­ Elgin, ll.) to supplement the fat and protein tains about 25-32% total solids, 5-12% fat, level of their milk replacer (Table 1). 14-19% crude protein, and 2-6% lactose In addition to a staple milk diet, many

180 ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY

Table 1. Calf feeding protocol provided by 18 different zoos and research facilites bottle raising moose calves. Facility Ingredient Feeding Schedule No. % No. Diet• ratio Age(weeks):vo1(l)lday:feedings/day fed raised Weaning date

1 CM:EM:EY" 2.5:1:1• 0-2:1.8:6,3-7:2:4,8-11:1.5:3, >11:0.9:1 7 86 late-Sep 2 CM:EM 2.0:1 1:2.2S:S, 2-3:2.S:S, 4-S:2. 7.S:S, 6-7:2.4:4, 8:1.8:3, 9-12:1.8:2 8 100 12weeks 3 CMR:CM l.S:l 0-12:2.5-6.0:5 depending upon age 60 mid-Aug 4 H20:EM:EY• 2.0:1:2" all ages:2.04.0:4 depending upon age 1S not provided S CM:EM 5.0:1 04:2.0:8, S-8:>2.0:6, 9-12:>2.0:4 28 15 by4months

6 EM:~O 1.0:1 1 :3~g. maximum of 2000ml:?, increase ratio to 2:1 >week 1 poor 40days 7 CMR 0-3:1.5:6, >3:2.4:5 14 36 late-Aug

8 EM:~O 1.0:1 all ages:llmglkg:S or 4 99 4months 9 CM:EM 1.0:1 0-3:2:6, 4-7:2.5:4, 8-12:reduced to 0 so 12wedcs 10 CM:EM 2.0:1 0-5:2.1:5,6-7:1.75:4,8-9,1.25:3, >9:0.3:2, 18 100 lOweeks 11 DM 04:2.8:3, S-8:2.85:2, 9-12:1.4:1 2 so mid-Aug 12 GM:CMR 1.0:1 all ages:3.2-4.0:4 3 100 12weeks 13 CM:EM• 2.5:1 0-8:2-6:4lamb replacier after 8 wks 100 12-16weeks 14 CM:EM 2.0:1 0-6:2.5:4,7-12:3.5-1.0:4, reduce to I liter by age 12wks 70 late-Aug 15 CMR 04:2.1:6, S-8:1.9:4, 8-12:1.2:4-1 16 81 early-Sep 16 CM:EM:EY• 1.0:1:4• 04:1.1:6, S-12:1.4:4, 13-16:1.9:4 90 4months 17 CMR 0-2:2.4-3.4:5,3-4:3.8-4.2:5,5:3.3:4,6-9:2.9:3, 10-14:1.8.0.9:2-1 39 59 late-Aug 18 CMR.r all ages:2S mglkg:8 2S 36 12-16 weeks

"eM-hole homoganized cow milk, EM=evaponted milk, CMR=cow milk replacier, ~<>=water, GM=, LMR=lama milk replacier, EY = egg yolk.

~e egg yolk was added to lliter whole cows milk mixed with 1 can evaporated milk (38Sml). "Two egg yolks were added to 1 can evaporated milk with 2 cans water. 'Thirty mls of was fed once per day. •Faur egg yolks were added to 1 pint can evaporated milk and 1 pint cow milk.

rc.seine was added to the milk to bring the crude fat level to 24%.

facilities indicated that they provided mineral calf attributed to white muscle disease. and vitamin supplements orally (mixture in Two facilities fed their calves probiotics the formula) or as an injectable. Both fat (A, (gastrointestinal tract microbial product) to D, E) and water soluble vitamins (B-complex) ensure a healthy development of lower gut were used. Injectable vitamins, although re­ flora. These products are relatively new, quiring additional time to administer, insured manufactured for domestic livestock, but may proper absorption, while those orally admin­ prove beneficial for rearing moose calves. istered may not be absorbed if the calf was Their function is to reduce dianbea; they also experiencing any digestive inflammation as­ enhance dry matter intake. sociated with dianbea. The amount of milk fed to calves depends Two facilities used injectable selenium to upon the size ofthe calf and the concentration prevent white muscle disease. We have also of nutrients in the milk itself. Most facilities used this at the MRC, where we have had at fed each young calf 2-2.5 liters/day (range least one documented fatality in an orphaned 1.1-3.8 Vday), in 5 or 6 feedings (range 3-8)

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MOOSE HUSBANDRY- SCHWAR1Z ALCES SUPPLEMENT 1 (1992)

spaced throughout the day (0600-2200 hrs) kept calves in close confinement for the first (fable 1). A few facilities indicated that they 2 weeks of life and then released them into a fed young calves (generally< 2 weeks of age) large pen. These calves were "called home" during the early-morning hours (0200 hr). at feeding times, but foraged on natural veg­ Three facilities fed bovine mixed etation during the remainder of the day. This with the fonnula the first week. technique reduced the need for an elaborate Milk volume increased and feedings per rearing pen and substantially reduced labor day decreased as calves aged. Generally milk cost to keep it clean. Calves reared this way volume fed to calves peaked when calves remained tractable because of their continued were 6-8 weeks of age, to a maximum (2.5-4 dependence upon bottle feeding. ]/day, range 2.0-6.0); the number of feedings Other useful suggestions included treat­ per day declined to 3-5 (range 2-6). Most ing the umbilicus of newly born calves with people weaned calves at 12 weeks of age iodine to reduce infections, isolation of sick (late-August) but one facility weaned at 10 animals, and regular replacement of bedding. weeks and several waited untillate-September Feeding Adults-Only 2 of26 (8%) facilities when calves were 16 weeks old. Weaning kept moose in pastures large enough to sustain was gradual with a reduction in milk volume them on natural browse. The remaining 24 and feedings/day. (92%) fed moose a fonnulated diet, and 21 Calf moose began sampling solid foods (88%) supplemented the ration with natural as early as 3 weeks of age, and readily con­ browse. In most cases, browse was harvested sumed solids by one month of age. Virtually and taken to the moose, although 4 facilities everyone raising calves offered browse (wil­ indicated that some natural browse and/or low, birch, aspen, fireweed), fonnulated ra­ aquatic plants were available within the hold­ tions, trace mineral salt, soil, and water. Many ingpens. indicated the sooner calves were switched Fonnulated rations were generally a mix­ from milk to solid foods the greater their ture of cereal grains (com, oats, barley) for chance of surviving. energy, soybean meal for protein, and Many people provided additional sugges­ woodfiber (sawdust) or hay as a fiber source. tions that warrant discussion Besides good They also contain vitamins, minerals, flavors, hygiene and immediate attention to diarrhea, and usually a product to enhance pelleting. which will be discussed in more detail later, Of 25 facilities providing the composi­ keen observation and "tender loving care" tion of their ration, 18 (72%) used sawdust were two suggestions most commonly listed. based diets, 4 (16%) used alfalfa (Medicago Moose calves responded to attention and were sativa) based diets, and 3 (12%) fed wood highly interactive with their caretaker. This fiber and grain. The most common diet listed interaction was the only substitute for the was the one we use at the MRC (Schwartz et mother-young bond and likely served to calm al. 1980, Schwartz et al. 1985). The diet the calf. During these interactions, caretaker contains an aspen byproduct (Table 2) as the noted changes in appetite, brightness of eyes, fiber source, is about 56-60% digestible, and general vigor, and possible looseness ofstools. has a crude protein content of 10-12%. Several people suggested that calves did Five facilities listed 3 modifications to best when raised outside; one person indi­ this diet Two facilities substituted sunflower cated that their calves did poorly when kept hulls for the aspen sawdust (Syroechkovsky indoors in a heated building. The best option eta/.1989:375). Chemically,sunflowerhulls was to rear calves in a large pen containing are similar to wood fiber (Anon. 1971), with adequate vegetation and food. One facility lignin being the main fiber type. This sun-

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;:' ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY flower hull diet was probably an adequate and sucrose. Sucrose is the energy source substitute, although neither facility fed it to replacing the com, barley and oats in the MRC moose for more than 4 years. Long-tenn diet. Sucrose is a readily digestible carbohy­ testing will detennine diet success. One facility drate. The manufacturer suggests that diets reduced the amount of com and increased the high in starch (energy source in cereal grains) concentration of soybean meal to raise the cause digestive problems in moose. They feel crude protein content to 14%. Two other that flora in the rumen of moose inadequately facilities substituted 10% alfalfa meal for part digest starch. This undigested starch then of the energy grains. Both facilities were passes into the lower gastrointestinal tract. small and kept only a few moose. Neither Fennentation of starch in the lower gut results facility experienced digestive problems in in chronic diarrllea (B. Sadler, pers. comm.). adult moose. No published data or controlled studies were Two facilities used a fonnulated ration available to support these claims. The com­ that contained about 30% dehydrated alfalfa pany is currently testing the diet at selected as the fiber source. Two facilities used a zoos. Purina also manufactures a breeding commercial brand of or deer ration, but diet (Mazuri Moose Breeder) which contains failed to provide me with the ingredient label. 16% crude protein and more digestible en- I assumed the source of fiber was alfalfa. One facility used the deer ration to supplement Table 2. Composition of the Moose Research Center ration. moose feeding in a large natural pasture. At one of the facilities, chronic digestive upset Ingredient % resulting in death was evident from necropsy corn, ground yellow reports. 30.0 Three facilities provided wood fiber along Sawdust 25.0 with concentrates. One facility fed oats, peas, Oats, ground 15.0 and poplar sawdust. A second fed hay helper Barley, ground 12.5 (product name and fonnula not provided) Cane molasses, dry 7.5 mixed with 10% birch sawdust. The third fed a 50:50 mixture of an aspen bark product Soybean meal (7 .4% nitrogen) 6.3 (80% aspen bark, 10% rolled oats, and 10% "Pelaid"• 1.3 rolled com, resulting in 3. 7% crude protein) Dicalcium phosphate 1.1 and commercial concentrate (equal amounts Sodium chloride 0.5 of com, oats, and barley plus molasses). One respondent listed laminitis as a problem in Vitamin A-D-Eb 0.3 adult bulls, but did not indicate if it was "Mycoban"c tr related to diet. At one facility they recently Trace minerals and flavorsd tr changed diets because of chronic diarrllea in adults and hoped the inclusion of wood fiber •"Pelaid" (Rhodera Inc., Ashland, Ohio) is a would cure the problem. pelleting enhance. A new diet currently being fed by one zoo bEach kg feed contained 5,000 IU Vitamin A, is the Mazuri Moose Maintenance Diet 13,000 ICU Vitamin 0 3 and 44 IU Vitamin E. (Purina, Saint Louis, Mo). The primary in­ c"Mycoban" (Van Waters and Rogers, Anchorage, gredient, wood flour, is aspen sawdust sized Alaska) inhibits mold growth, tr =0.025%. to pass through a pelleting machine. The diet 4Added trace minerals and flavors (mg/kg) Zn, contains 11% crude protein. It differs from 55.2;Fe,30.5;Mg,22.0;Mn, 16.8;Cu, 7.6;I, 1.3: the MRC diet because it contains alfalfa meal and anise, 125.

183 T i MOOSE HUSBANDRY-SCHWARTZ ALCES SUPPLEMENT 1 (1992)

ergy. If these diets prove successful, avail­ made available, particularly if browse con­ ability of a fonnulated moose ration will im­ tains green leaves. During summer when prove. consumption rates are high, loosely fonned Dr. C. T. Robbins at WashingtOn State stools resembling "cow pies" are common. University has also fonnulated a starch free Healthy animals produce typical moose pel­ test ration for moose (Table 3). He has suc­ lets during winter. cessfully fed this diet for more than 2 years. A Weaned calves consume about 1.5-2.5% bull consuming the MRC ration with indica­ of their body weight/day. Calves will main­ tions of diarrllea produced nonnal pelleted tain a relatively high rate of intake during feces when switched to this diet. Controlled winter and continue to gain weight (Schwartz experiments evaluating starch digestion in et al. 1984, Schwartz et al. 1987b). moose seem warranted. Since this survey was conducted (1990) Table 3. Composition of the Washington State and the time of printing (1992), 3 facilities University moose mtion. have tested the Mazuri ration. Two of three facilities are pleased with the diet One of­ Ingredient % fered both the Mazuri and MRC diets in Sawdust 22.5 combination and noted that the animals Canolameal 20.0 switched freely between the two, consuming Beet pulp 18.8 more of one diet during certain times, but less during others. The third has experienced Alfalfa meal 10.0 some digestive problems with one animal, but Molasses (dry) 8.0 the cause was uncertain. Sucrose 7.5 Besides offering fonnulated diets and Wheatbmn 5.0 browse, several facilities (36%) regularly feed produce (carrots, apples, bananas, lettuce, Bone meal 5.0 cabbage) and bread. Many people felt these Animal fat 0.75 foods were enjoyed by the animals and served Tmce mineral salt• 0.75 as a dietary treat. Almost 20%, supplemented Lime 0.50 moose diets with hay, mainly alfalfa or an alfalfa-bromemix. One facility supplemented Dicalcium phosphate 0.05 with grass hay and another with clover hay. Vitamin premixb 0.25 No one fed grass hay alone or as the major Vitamin A premix 0.225 dietary item. Se premix (0.02% Se) 0.150 Food Consumption Rates-Our experience Vitamin E premix (500 IU/g) 0.070 at the MRC indicates that food consumption will vary with season and sex ofmoose. Both Vitamin K premix (0.14 g/g) 0.005 males and females consume about 2.5-3% of their body weight of dry feed in summer. "Tmce mineral salt: 96-96.5% NaCl, 0.20% Mn, They reduce this level to 0.5-1.0% of body 0.35% Zn, ).20% Fe, 0.15% Mg, 0.03% Cu, 0.007% I, and 0.005% Co. weight in winter (Schwartz et al. 1984). Re­ duced consumption in winter is voluntary and bVitamin premix: 400,000 IU Mb, 30,000 IU Dj no reason for alann. Bulls do not eat during lb, 2,000 IU E/lb, 360 mg B/lb. 1,200 mg niacin/ the rut for as long as 3 weeks; this fasting is lb, 1,200 mg d-panothenate/lb, 45,000 mg choline/lb. 1.8 mg B flb, 90 mg menadione so­ nonnal (Schwartz et al. 1984). Consumption 1 dium bisulfite/lb. 100 mg folic acid/lb, 9 mg d­ rate of a fonnulated diet declines as browse is biotin/lb, and 4,540 mg ethoxyquin/lb. 184 ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY

Bulls reach peak body weight in late­ (77%) breed as yearlings. Augustorearly-Septemberjust before rut and Primiparous females consistently pro­ velvet shedding. They can lose up to 20% of duced a single calf, although 2 sets of twins this weight during the rut, but generally regain were reported. At the MRC, 1 yearling cow body condition in November and early De­ out of 10 produced twin calves. Facilities cember. Overwinterweightloss in bulls ranges indicated that adult females produced either 1 from 7% to 23% depending upon conditions. (65%), 2 (10%), or 1-2 (25%) calves. Few Cows reach peak body weight in mid-winter, facilities provided reproductive histories of much later than males, and minimum weights individual females. At the MRC, of 31 births occur after parturition. Average weight loss by mature cows, 58% were twin and 42% ranges from 15-19%(Schwartzetal.l987b). were single births; we have never had triplets. This past winter, we experienced a de­ Our overall productivity of calves from all cline in intake associated with a change in the females averaged 1.5 calves/birth. diet constituents of the feed. Our feed manu­ No respondent felt they had major repro­ facturer replaced dry cane molasses with liq­ ductive problems. Minor problems included: uid molasses. Although the change appeared still births (n = 12), adult cows not producing minor, we experienced more than a 50% de­ calves (n = 3), dystocia, difficulty birthing, or cline in food intake; several moose refused to breached births (n =4), spontaneous abortions eat this new diet We are unsure why liquid (n = 2), defonned calves (n = 2), retained molasses made the ration less palatable, but placenta (n = 2), uterine inertia (n = 1), and suspect ingredient quality, concentration, odor, failure to come into heat (n = 1). In addition, and possibly discriminating individuals. This several people (38%) indicated that their cows incident does suggest caution when altering gave birth to small, generally frail calves that diet make-up. died shortly after birth (n = 25). Several people cautioned about the potential danger to Breeding caretakers from rutting males. Fourteen facilities breed moose. Twelve facilities breed cows as soon as they reach Herd Health Problems sexual maturity, whereas 2 did not breed year­ Calves-Diarrhea is a common problem ex­ ling. At the MRC, annual calf production perienced by most facilities raising calves. over at least a 10 year period for each cow Loose stools seem to be inevitable with bot­ averaged 1.5 calves for females (n = 4) that tle-raised calves and should not be cause for were first bred as yearlings and 1.9 calves for concern unless they are severe. Severe dianbea females (n = 3) first bred as 2-year-olds. involves loose, watery feces, which is often These data suggest calf production over the foamy and released involuntarily. Chronic breeding life of a cow may improve by not long-tenn diarrhea ultimately results in weight breeding females as yearlings; additional re­ loss, dehydration, and in severe cases, death. search is needed. A number of facilities treat dianhea with Sexual maturity and age offirst reproduc­ medications, primarily Kaopectate (Upjohn tion in moose are directly related to body size Co., Kalamazoo, MI 49001) or some other and nutrition (Simkin 1974). In captivity, brand of anti-dianhea medicine available at females first breed at 1.5 (57%), 2.5 (29%) or the local phannacy. A few facilities also use 3.5 (14%) years-of-age. This high rate of sulfa drugs. A common practice to help reproduction in yearlings indicates the high alleviate dianhea is to eliminate milk from the plane of nutrition of captive reared moose. At diet for one to several feedings and substitute the MRC, we have had 10 of 13 females an electrolyte solution that contains glucose.

185 MOOSE HUSBANDRY-SCHWARTZ ALCES SUPPLEMENT 1 (1992)

A second problem that appears related to provided necropsy records and/or listed the bottle-raising calves is laminitis or foundering. cause of death for animals dying at their Foundering in young ruminants is caused by facility. I reviewed these reports and catego­ carbohydrate overload that results in degen­ rized deaths into infectious and noninfectious eration and inflammatory changes in the lami­ categories that I subdivided further (Table 4 ). nae of the hoof (Jubb et al. 1985). Extreme Infectious diseases accounted for 32% of changes in diet quality and volume cause all deaths. Bacterial and viral agents were laminitis in bottle-raised moose calves. Calves responsible for 69% of infectious diseases. can live with the affliction, but as they mature Moose died with pneumonia, systemic infec­ their weight, the leverage force placed on the tions, gastroenteritis, and malignant catarrhal toes, and the pulling forces of certain tendons fever. Malignant catarrhal fever is a viral results in poor alignment of the foot and leg infection that is lethal to moose. Domestic bones. There is a general elongation of the sheep and other members of the sheep and hoof and varying degrees of pain and diffi­ goat family (Bovidae) carry the disease. In 2 culty associated with walking. cases, Ibex (Capra aegagrus) and domestic Polyarthritis and meningitis are two ail­ sheep were the suspected reservoir. ments that afflict bottle raised calves. Both The other form of infectious disease re­ canresultfromageneralsepticemiaassociated sponsible for 10% of the deaths (31% infec­ with microbial infections. Invasion may oc­ tious deaths) was parasitic meningeal worm, cur through a wound or in very young calves whip worm, thread worm, and hydatid cysts through the umbilicus (Timmermann and (Table 4). Surprisingly meningeal worm Lancaster 1978). (Parelaphostrongylus tenuis) was only re­ Adults-Several of the reporting facilities ported once in spite of the very common

Table 4. Reported cause of death for 45 adult (age> calf) moose held in captivity in zpos, game farms, and research facilities in North America.

Cause of death n % Comments Infectious 32 Bacterial or viral 22 Gastroenteritis 2 4 Salmonella spp, Escherica coli Malignant catarrhal fever 3 7 carried by sheep and Pneumonia 5 11 may have been secondary agent Parasitic 10 Brain worm 1 2 Parelaphostrongylus tenuis Whip worms 2 4 Trichuris spp. Thread worms 1 2 Trichostrongylus spp. Hydatid cyst 1 2 Echinococcus granulosis, pneumonia Noninfectious 67 Man-caused 7 16 Complications during immobilization Nutritional 6 13 poor body condition, teeth, chronic wasting Digestive upset 14 31 chronic diarehha, gut ulceration, torsion, Arthritis and laminitis 3 7 predisposed to pneumonia or poor nutrition

186 ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY

practice of mixing hoofed stock, particularly degree of health problems, especially facili­ white-tailed deer (Odocoileus viriginianus) ties with small enclosures. andwapiti(Cervuselaphus)withmoose. Both As with calves, laminitis or foundering, carry P tenuis, which is lethal to moose (Lan­ predisposed a small number (7%) of animals caster 1987). to problems and death. Arthritis was less The low percentage of deaths caused by common, probably because few moose live parasites may in part be due to control meas­ long enough to develop the problem. ures. Most facilities routinely (2-6 times/ Common health problems not responsi­ year) examine feces for parasites. Periodic ble for deaths included winter tick infestation treatment with Ivennectin was a common (Dermacentor albipictus), biting insects, over­ practice. grown hoofs and dewclaws, and injuries to Noninfectious diseases and accidents were antlers. Pesticides controlled insects and ticks. responsible for the greatest percentage ( 67%) Longevity-The average life-span ofa moose of all reported deaths in adults. Drug related kept in captivity listed by those facilities re­ deaths were responsible for 16% ofthe known sponding ranged from 3-8 years, with the mortality, mainly from aspiration pneumo­ majority reporting 4-6 years. The longest nia, but also from capture myopathy. lived animals were reported at 8-13 years of The single greatest cause of death listed age, with the majority listed at 10-11. No was nutritional and digestive upset resulting facility reported keeping a moose over 13 from chronic diarrhea. Nutritional problems years old. This is much younger than similar ranged from starvation resulting from reports from wild populations where moose malocclusions and/or badly worn teeth (n = have been known to reach the age of 16-20 3), injury to the tongue (n = 1), and improper (Peterson 1955). feeding (n =2). Both old and young animals Several zoos provided taxon reports which had tooth problems. Other respondants re­ list date of death for moose kept in captivity. ported worn teeth not associated with death. It Using these data, I constructed a survival appears that tooth problems are common to curve (Fig. 1) which depicted life expectancy moose in captivity. in captivity. Estimates were minimum since Digestive upset, often resulting from animals still living were not included. chronic diarrhea was the single most common The single greatest age class suffering cause of death (31% ). It was difficult to high rates of mortality was calves. Almost detennine why moose died from this ailment, 70% of all calves raised in captivity died be­ but I suspect it was dietary. Animals dying fore their first birthday. This estimate includes from long-tenn chronic dianhea and loss of body condition, generally were fed for several years on diets containing grass hays or a fonnulated ration containing crop residues or hay. However, 2 facilities that fed both hay­ based and sawdust-based rations reported z 0., 0 similar problems with both diets. One facility ~ o... held moose in captivity for many years(> 30), "'a.' 0.3 li! 0.2 <>. and the other held moose for 13 years. One 0.1

facility fed browse regularly, the other rarely. 0 ~o---1--+2--+3--+,---,--~,---7---.---.~1o· Not all problems are solely dietary. There AGE IN YEARS seemed to be some relationship between the Fig. 1. Proportion of moose kept in captivity sur­ number of years a facility kept moose and the viving by age class in North American zoos and research facilities. 187 MOOSE HUSBANDRY-SCHWARTZ ALCES SUPPLEMENT 1 (1992) all sources of mortality. Since some facilities with post spaced 3.0-4.6 m apart. Fencing commonly attempt to raise orphaned calves, costs at least $8,000/km just for materials low survival is expected. Orphaned calves are (Renecker et al. 1987). Holding facilities often weak, dehydrated, injured, and may not must be large with a stocking rate of about 1 have eaten for several days. Chances of moose/U.S ha when the animals are fed sup­ survival are low. plemental rations. Few facilities keep more than 5 moose together. Due to the solitary Compatibility of Other Species nature of moose, keeping large numbers of Over half the facilities reporting indi­ animals togethermay be detrimental, although cated that they kept other species with their additional studies on social stress are war­ moose. Ungulate species included white ranted. tailed-deer (n =6), mule or black -tailed deer In northern climates, moose appear to do (0. hemionus) (n = 5), wapiti (n = 6), caribou well with minimal shelter. Dense stands of or (Rangifer tarandus) (n = 3), trees or a 3-sided shelter provide adequate pronghorn (Antilocapra americana) (n =2), protection from winter -storms and wind. bison (Bison bison) (n = 1), During summer, temperatures exceeding 27"C (Ovis canadensis) (n = 1), can cause thennal stress and adequate cooling (Oreamnos americanus) (n = 1), fallow deer facilities are desirable. Trees and shelters (Dama dama) (n = 1), and (Ovibos work in northern climates, but ponds, streams, moschatus) (n = 1). In addition to other or sprinkler systems are more appropriate in hoofed stock, three facilities kept waterfowl hot climates. and gallinaceous birds, and one facility kept Raising calves is a difficult task. Chronic black bear with the moose. The facility keep­ diarrhea is the most common cause ofdeath in ing black bears did experience some calf bottle-raised orphans. Milk replacer made predation and was forced to separate the bears with a mixture of whole cow milk and from the moose during calving season. No evaporated milk or a dairy milk replacer is the respondent indicated any behavioral problems most common fonnulas used. Success with or fighting between the mixed hoof stock and each fonnula is difficult to detennine, but it moose. Several people indicated moose were appears that the cow milk and evaporated very compatible with other species. milk mixed diet is slightly superior. Diet evaluation requires better record keeping. SUMMARY Supplements to increase the fat and protein The number of facilities keeping moose content of diets may be useful but their needs in North America has increased in recent and success require further evaluation. years. Commercial game fanning is a growing Moose are a sensitive species in captivity. industry (Hudson et al. 1989). Although bi­ Meeting feed requirements appear to be the son, reindeer, and wapiti are the most profit­ single greatest hurdle. The development of a able ungulates, moose have served in a lim­ relatively new ration that contains wood fiber ited way to diversify these grazing systems in the fonn of aspen sawdust apparently meets (Renecker et al. 1987). Currently, moose are both nutritional needs and physical diet struc­ raised for display or sale as breeding stock. ture. Its development allows for moose to be Because moose are difficult to keep and ex­ kept in captivity more easily than feeding pensive to feed (Schwartz et al. 1985), they them browse (Ellis 1987). Small amounts of are not raised for meat or milk production. grass hay fed to moose for short periods cause no apparent problems. However, necropsy Moose are eas~ly kept inside standard game fencing that is a minimum of2.1 m high reports suggested that long-tenn feeding of

188 ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY grass hay rations causes digestive upset and (Goplen et al. 1980). Secondary plant chronic diarrhea which can result in death. metabolites may serve a useful purpose in the Additional research is required. digestive system of moose. The compounds When I first developed the MRC ration may be responsible or aid in the maintenance (1978), I was convinced that many of the of specific environments needed by lower gut digestive problems in captive moose were flora. They may prevent infections or invasion dietary. Improper diet seemed to be the cause of certain organisms. In many cases, when a of chronic wasting in moose and facilities "sick" moose is provided natural browse the which kept moose for many years had more problem clears. Something in natural foods is problems. Itis unclear ifthe problems resulted lacking in our synthetic rations. Sawdust from: (1) increased exposure and/or accu­ based rations have partially met the challenge mulation of some infectious agent, (2) an of matching the synthetic diet with natural increased probability of experiencing prob­ foods, and removing starch may be another. lems with time, and/or (3) it requires time for However, we must continue to investigate chronic wasting to develop even in improperly other components that might be "required" by fed moose. the animal in the non-classical sense. Ad­ If wasting is dietary, we must attempt to vances in the field of moose nutrition will be match our synthetic diets as closely as possible enhanced with continued cooperation and to the natural diets of moose. Thus we must coordination among all the facilities keeping consider more than simply classically defined moose in North America. "required nutrients" (e.g., protein, energy, Live expectancy of moose in captivity is macro and micro elements, or vitamins). less than in the wild. In North America it is Matching the natural diet means providing still difficult to keep moose under strictly compounds which are possibly "essential" for confined conditions. Mortality in captivity reasons other than they are required in a seems to result from disease and nutritional metabolic sense. For example, moose are problems. Although our understanding of the selective browsers and consume large quan­ nutritional requirements of moose has in­ tities of secondary plant compounds (tannins, creased in recent years, there is still much to phenolics, and others). Evolutionarily, the learn. Chronic diarrhea has been a major moose has adapted to handle these chemicals. problem in the past and still persists today. Classically, we think of secondary plant com­ Proper husbandry minimizes infectious dis­ pounds as "bad" for the animal. They are ease. Simple practices, such as not mixing compounds which the plant has produced to moose with other ungulates that carry fatal prevent herbivory. However, since moose disease or parasites seem logical. Frequent evolved to accommodate such ''toxins" ,moose screening of feces to detect parasites and may "requires" or "needs" them in a non appropriate treatment with medication is nec­ classical sense. Tannins in diets ofleaf-eating essary. Laminitis, arthritis, and hoof insects, for example, bind with certain viral overgrowth can cause chronic problems. A agents and prevent infection. Insects eating hard substrate (near food or water source so diets with tannin had lower mortality rates they cross it daily) and periodic hoof trim­ associated with a polyhedrosis virus than in­ ming prevents hoof overgrowth. sects consuming similar foods without these Moose reproduction in captivity is simi­ tannins (Keating et al. 1988). Domestic live­ lar to that in the wild. Most females reach stock consuming legumes which contain sexual maturity as yearlings, but to maximize tannin seldom develop bloat, while those con­ calf production they should probably not be suming legumes without tannin get bloat bred until 2.5 years of age. Twins are com-

189 T

MOOSE HUSBANDRY-SCHWARTZ ALCES SUPPLEMENT 1 (1992)

mon from adult cows although single calves panmentoflnland Fisheries & Wildlife, Gray, are also produced. Calf production averages Maine; Fred Servello, Depanment of Wild­ 1.5-1.8 calves/year over the life of a female. life, University of Maine, Orono, Maine; Calf survival varies, but small weak calves Robyn Barbiers, Detroit Zoo, Detroit, Michi­ that die shortly afterbirth are common. Moose gan,NickReindl,MinnesotaZoo, Apple Val­ occasionally suffer from dystocia, abortion, ley, Minnesota; Don Winstel, Columbus Zoo, failure to breed, and retained placenta. Al­ Columbus, Ohio; Dave Ellis, Northwestrek, though up to 5 generations of moose have Eatonville, Washington; Charles Robbins, beensuccessfullymaintainedincaptivity,most Depanment of Natural Resources, Washing­ facilities routinely supplement their stock with ton State University, Pullman, Washington; animals from the wild. ElizabethFrank,MilwaukeeZoo,Milwaukee, When I set out to consolidate what has Wisconsin; Huey Dawnson, Sybille Wildlife been learned about moose husbandry in North Research Unit, Wyoming Game and Fish, America, I had no idea what kind of responses Wheatland, Wyoming; LyleRenecker,School I would receive. I was very pleased that of Agriculture, University of Alaska, virtuallyeveryonekeepingmooseparticipated. Fairbanks, Alaska; Wayne Ray, Anchorage If we all worlc. together, we can continue to Zoo, Anchorage, Alaska; Murry Lankester improve our understanding of moose require­ and Steve Dudzinski, Depanment ofBiology, ments in captivity. Facilities keeping moose Lakehead University, Thunder Bay, Ontario; should make every effon to keep accurate Board ofManagement, Metropolitan herd records. Only with a clear understanding Zoo; and Jacques Prescott, Jardin Zoologique of husbandry problems will we ultimately du , Charlesborug, Quebec. I also succeed in our efforts to successfully main­ thank Kris Hundenmark, Melody Schwartz, tain this unique deer species in captivity. Karl Schneider, C.T. Robbins, and Sid Morgan for reviewing the manuscript and David C. ACKNOWLEDGEMENTS Johnson for summarizing much ofour histori­ I would like to thank all those people who cal data from the Moose Research Center. participated in the husbandry survey: B. L. This study is a contribution of Federal Pendelton, Forestry Farm Park & Zoo, Aid in Wildlife Restoration, Project W-23-4. Saskatoon, ; Donna McAllister, Elk Valley Ranches, Kitscoty Albena; Bob REFERENCES and Lynn Stewan, Terrestrial & Aquatic ADDISON, E. M., R. F. MACLAUGHLIN, Environmentals, Melville, Saskatchewan; and D. J. H. FRASER. 1983. Raising Peter Kalden, Davis Point Game Farm, St. moose calves in Ontario. Alces 18:246- Martin, ; Dean Treichel and David 270. Leeb, Valley Zoo, Edmonton Albena; Pam ADDISON,E.DODDS,D.F. 1959. Feeding McDougall, Zoo, Calgary, Albena; and growth of a captive moose calf. J. Mac Pitcher, Salmonier Nature Park, Wildl. Manage. 23:231-232. Holyrood, ; Pierre Poirier, ANON. 1971. Atlas of nutritional data on , Granby, Quebec; Sandra United States and Canadian feeds. Na­ Bergeron, Zoo de St-Felicien, St. Felicien, tional Academy ofSciences, Washington, Quebec; Hugh Oakes, VancouverGameFann, D.C. 642pp. Algergrove, ; Bruce Dougan, ELliS, D. E. 1987. Moose nutrition: a Magic Hill Zoo, Moncton, New Brunswick; successful longterm captive diet. Proc. Ed Addison, Ministry of Natural Resources, N. Am. Assoc. of Zoological Parlc.s and Maple, Ontario; David Wilbur, Maine De- Aquariums Conf. 190 ALCES SUPPLEMENT 1 (1992) SCHWARTZ- MOOSE HUSBANDRY

FRANZMANN, A. W., P. D. ARNESON, Congr. Game Bioi. 14:425-429. and D. E. ULLREY. 1975. Composition RENECKER, L. A. 1987. The composition ofmilk from Alaskan moose in relation to of moose milk following a late parturi­ other North American wild ruminants. J. tion. Acta Theriologica 32:129-133. ~.Anim.~ed.6:12-14. ___,andR.J.HUDSON. 1986. Seasonal ___,C. C. SCHWARTZ, and D. C. energy expenditure and thennoregulatory JOHNSON. ~onitoring status (condi­ response of moose. Can. J. Zool. 64:322- tion, nutrition, health) ofmoose via blood. 327. Swedish Wildt. Res. Suppl1:281-288. ___, ,andG. W.LYNCH. 1987. GOPLEN, B.P., R. E. HOWARTH, S. K. ~oose husbandry in Albena, Canada. SARKAR, and K. LESINS. 1980. A Swedish Wildt. Res. Suppl. 1:775-780. search for condensed tannins in annual ROBBINS, C. T. 1983. Wildlife feeding and and perennial species of M edicago, nutrition. Academic Press, New Yorlc, Trigonella, and Onobrychis. Crop Sci­ N.Y. 343pp. • II ence. 20:801-804. SCHWARTZ,C.C.,W.L.REGELIN,andA. HJEUORD, 0. 1987. Research on moose W.FRANZMANN. 1980. Afonnulated nutrition in the Nordic countries. Swed­ ration for captive moose. Proc. N. Am. ish Wildl. Res. Suppl1:389-398. ~oose Conf. Worlcshop 16:82-105. HUDSON, R. J., K. R. DREW, AND L. ~. ___, and . 1984. BASKIN. 1989. Wildlife production Seasonal dynamics of food intake in systems. Economic utilisation of wild moose. Alces 20:223-244. ungulates. Cambridge Univ. Press, ___, and . 1985. Cambridge. 469pp. Suitability of a fonnulated ration for JUBB, K. V. F., P. C. KENNEDY and N. moose. J. Wildl. ~anage. 49:137-141. ___., and ~. PA~R. 1985. Pathologyofdomestic animals. Academic Press, Inc. New York. HUBBERT. 1987a. Nutritional energetics of moose. Swedish Wildl. KEATING, S. T., W.G. YENDOL, and J. C. Res. Suppl. 1:265-280. SCHULTZ. 1988. Relationship between ___, and . 1987b. susceptibility of gypsy moth larvae Seasonal weight dynamics of moose. (Lepidoptera: Lymantriidae) to a Swedish Wildl. Res. Suppl. 1:301-310. baculovirus and host plant foliage con­ SI~IN. D. W. 1974. Reproduction and stituents. Environ. Entomol. 17:952-958. productivity ofmoose. Naturaliste can. LANKESTER.~. W. 1987. Pests,parasites 101:517-526. and diseases of moose (Alces alces) in SPEIDEL, G. 1965. Care and nutrition of North America. Swedish Wildl. Res. moose (Alces alces americana) in cap­ Suppl. 1:461-489. tivity. Inter.~ Yearbook 6:88-90. ~ARKGREN, G. 1966. A study of hand­ SYROECHKOVSKY, E. E., E. V. reared moose calves. Viltrevy 4:1-42. ROGACHEV A, and L.A. RENECKER. PETERSON, R. L. 1955. North American 1989. ~oose husbandry. Pages 369-386, moose. Univ. Toronto press, Toronto in R. J. Hudson, K. R. Drew, and L. ~. Canada. 280pp. Baskin, eds. Wildlife production systems, REGELIN, W.L.,C.C.SCHWARTZ,andA. economic utilization of wild ungulates. W. FRANZMANN. 1979. Raising, Cambridge Univ. Press, Cambridge. training and maintaining moose (Alces TI~~ER~ANN, H. R., and ~. W. alces) for nutritional studies. Proc. Int. LANKESTER. 1978. Joint disease in

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