1¾«˜Z¾ĖĖĻŠŠCUBSFOOD.STYTECŅ@ŠpĢĖĪµHuber et al. -  - Send proof to: September 28, 1992 Djuro Huber Biology Department Veterinary Faculty Heinzelova 55 41000 Zagreb Republic of Croatia Phone 041/290-141 Fax 041/214-697 FOOD INTAKE AND MASS GAIN OF HAND-REARED BROWN BEAR CUBS Djuro Huber 1, Ignac Kulier 2, Andrea Poljak 1, and Branka Dev~i}-Kuhar 1 1 Veterinary Faculty, Heinzelova 55, 41000 Zagreb, Croatia 2 Pliva, Istra`iva~ki institut, Planinska bb, 41000 Zagreb, Croatia RUNNING HEAD: Growth of hand-reared brown bear cubs ABSTRACT Five orphaned European brown bear cubs (Ursus arctos) from three litters were hand-reared since the ages of 1 or 4 months and masses 1.7-2.8 kg. The growth rate was monitored with special reference to the food used. Over the period of 3 years 6 different feed formulas were used. The first four were bottle nursed till the average age of 133 days. The conversion of all feed formulas in the first 10 months ranged 3.5-32.0 g of food for g body mass (or 38.1-192.6 kJ/g body mass), and was influenced mostly by the food composition. The high content of fat showed as good, and high content of carbohydrates as bad indicator. The feed formula 3, with 12.0% proteins, 23.9% fats and only 0.2% carbohydrates, was a simulation of bear milk and showed the best results. The feed formula 6 (bread, fruits and meat) was used from ages of 7 to 35 months and over this period the conversion gradually decreased 3.8 times as a consequence of maturing process. Hand-reared cubs were 1.3-2.7 times larger than 17 measured wild cubs at matching ages, which is explained by limitations imposed to the mother by hibernation and mainly herbivorous diet. Key words: Ursus arctos, food conversion, growth INTRODUCTION Bears are born very small (<1% of mothers mass) and helpless in the middle of winter when no food, even for the mother, is available. Hence, the mother produces very concentrated milk (Jenness et al., 1972). In 3-4 months (January through April/May), on this exclusive diet, cubs have to grow big and strong enough to be able to leave the den and follow the mother in spring foraging. With the decreasing dependency the mother nurses her cubs at least 1.5 years. The food intake of cubs and early growth of European brown bears (Ursus arctos) have not been studied. In this respect much more is known about American black bears (U. americanus). Hock (1966) compared caloric intake and mass gain in two American black bears since the age of 21 days. Other aspects of postnatal growth of the same species were described by Rausch (1961), Butterworth (1969) and Hulley (1976). The composition of milk of different species of bears was studied by Jenness et al. (1972). Oftedal and Gittleman (1989) compared patterns of energy output during reproduction in all terrestrial carnivores. We monitored the growth rate, with special reference to the food used, in five orphaned European brown bear cubs from the ages of 1 or 4 months till the ages of up to 4 years. The influence of the degree of similarity of different food mixtures to the bear milk on conversion was analyzed, and masses of hand-reared and wild cubs were compared. MATERIALS AND METHODS The five studied orphaned European brown bear cubs belonged to three litters born in Gorski Kotar, Croatia and all were females. Cubs marked 1 and 2 belonged to litter A, 3 and 4 to litter B, and 5 to litter C (Table 1). The litter A had a third cub which died within 24 hours after coming to our hands. Post mortem of this cub revealed pulmonary oedema and emphysema with large milk curd obstructing the stomach. Occasional data of 17 other cubs nursed by their mothers were used for comparisons (original data and Frkovi}, 1983). The estimations of birth dates (Table 1) were made by an extrapolation of daily mass gains back to the expected birth mass (Hock, 1966). The state of physiological features like healing of the umbilicus, hair growth and eyes opening was also considered. In all later calculations of common age we used 12 January as the mean date of birth for all cubs. The cubs were given to us, at given dates (Table 1), for hand-rearing after they were found out of den with no mother around (litter A), after the mother was accidently chased out of den and did not return within 48 hours (B), or the cub wandered alone into the village (C). Over the monitored period the cubs were fed with 6 different feed formulas. They were individually bottle-nursed the first 4 mixtures and all consumed quantities were recorded. Later the cubs were eating solid food all together and the total consumed amount was divided equally to each bear. The food intake was recorded as the mass of prepared meal with the moisture included. The nutritional value (gross energy) of each formula was obtained by computer analysis. The first two mixtures were based on artificial formulas for human infants. The feed formula 3 was prepared by the second author as the simulation of bear milk. Periodical measurements of mass and 39 body sizes gained a total of 58 sets of bear measurements or the mean of 11.6 (range 3-18) per bear. Only data on body mass (Table 2) and total length were considered in this paper. After the age of three months the cubs had to be chemically immobilized (Ketamine-Ksilazine mixture) to take the measurements. Butterworth (1969) reported that he had to discontinue the measurements of the American black bear cub at the age of 102 days because the animal was difficult to handle. RESULTS The obtained results are presented by the sequence of the different feed formulas used. Considered are the compositions (Table 3), consumed amounts and conversion rates of all used foods (Table 4). The feed formula 1 was based on the daily combinations of the human baby feed formulas following the manufacturer's recommendations: A) formula for low mass infants (9.5% volume over the period), B) for healthy infants (58.1%), and C) antydiarrhea formula (29.5%). This food was used for the first 15 days for cubs 1 and 2 in their estimated ages from 29 to 43 days. Both cubs accepted it but had constant diarrhea. The cub 2 had pneumonia since it came to our hands and was under antibiotic treatment for 6 days. The average daily consumption of 508 g of prepared meal (116 g of dry food mass) distributed in an average of 6.0 meals (range 5-8) gained 54 g of body mass with conversion of 9.4 g food or 40.9 kJ per g mass gain. In the ad libitum (unrestricted) feeding of the both cubs, the cub 1 consumed 27% more than cub 2, her daily mass gain was 2.3 times bigger, and the conversion was 1.4 times better. The 2nd mixture contained formula for human babies over 1 month old (44.0%) and antydiarrhea formula (10.6%) with addition of egg yolks (17.6%) and cream with 30% fat (27.8%). It was used for 38 days for 4 cubs (1, 2, 3 and 4) in the ages from 44 to 81 days. The mean daily consumption of 707 g of prepared food, in 5.2 meals (range 4-8), gained 120 g of body mass with conversion of 5.9 g food or 46.8 kJ for g body mass. The cubs 1 and 2 were feed ad libitum and consumed 1.53 time more than cubs 3 and 4 whose meals were rationalized. The mass gain in the first two cubs was 2.4 times greater and the conversion 1.6 times better. At the age of 73 days cubs could drink water and 2 days later ate some food from the dish. The 3rd feed formula was a replica of the basic composition of the brown bear milk: in the dry mass it contained 63.6% fats, 31.8% proteins and 0.6% lactose. It was fed to 4 cubs as an exclusive diet for 33 days (ages 82-114 days). The average consumption of 649 g of prepared food in 3.4 (range 2-5) meals resulted with 188 g of daily gain with conversion of 3.5 g of food or 38.1 kJ for g body mass. Again cubs 1 and 2 consumed 1.97 times more than cubs 3 and 4 and gained 1.41 times more, but the conversion was only 0.67 of the latter two. The feed formula 4 was a combination of bear milk simulation (i. e. feed formula 3; 55.4%) and the full cow milk (44.6%). It was used for all 5 cubs for 19 days (estimated ages 115-133 days). The mean consumption of 759 g, in constant 3 meals, gained 101 g of body mass with conversion of 7.5 g or 54.8 kJ per g mass. The cubs 1 and 2 consumed 1.5 times more but gained only 0.78 compared to cubs 3 and 4. Also the conversion was only 0.52 of the one in the latter two cubs. The cub 5 consumed daily 729 g and recorded the high 138 g of day gain with the best conversion of 5.3 g or 38.6 kJ for g body mass. The feed formula 5 was a mixture of cow milk (66%) and solid food, i. e. hard-boiled corn mush and wheat noodles (paste) (17% each). The 5 cubs consumed every day 6000 g of this food from the central feeder, or an average of 1200 g each, for 51 days (estimated ages 135-185 days). The mean mass gain was 101 g with the conversion of 11.9 g or 60.1 kJ for g mass gain. At the average age of 186 days all 5 cubs were released to nature. However, cubs 1 and 2 after 17 days, and cub 3 after 47 days returned to people and had to be retaken for captive holding. While feeding on natural food cub 3 gained a mean of 85.1 g/day. Cub 4 disappeared and cub 5 was found dead after being struck by a vehicle. Since the ages of 222 days (cubs 1 and 2) and 233 days (cub 3) till the ages of 1046 days (cub 3) and 1387 days (cubs 1 and 2) cubs were kept in the Zoo of Zagreb and were fed the mixture 6 which contained bread (31%), apples or other fruits (64%) and beef meat and fat (5%) in the total amount of 5150 g per cub per day. The mean daily gain for 3 cubs (or 2 after 824 days) over 1165 days was 71 g with conversion of 74 g or 451 kJ of food for g body mass. However, in the 1st 84 days (est. ages 222-305 days) of feeding with the same amounts of this mixture the mean daily gain was 161 g and conversion 32 g or 193 kJ per g gain. In the next 379 days (est. ages 306-684 days) the daily mass gain was 100 g and conversion 52 g or 312 kJ per g gain. During the last 703 monitored days (est. ages 685-1387 days) the gain was 46 g and conversion 112 g or 677 kJ of food per g gain. The day gains during the use of the first 3 feed formulas, or over the ages from 29 to 114 days, cubs would double their body mass in the mean of 32 days (range 29-39). The day gains with feed formulas 4, 5, and the first 84 days of formula 6 (ages 115-305 days) permitted the body mass to double in a mean of 135 days (range 128-143). In the next 379 days of consumption of the 6th feed formula, 375 day gains were necessary to double the body mass. In the last 703 days with the same food, 1755 day gains would be necessary to double the body mass (Table 4). Seventeen wild bear cubs nursed by their mothers were measured in ages from 87 to 1243 days. They showed great variations of body masses but were 1.3-2.7 times smaller than our hand-reared ones at all comparable ages (Table 2 and Fig. 1). The body length of mother-nursed cubs was also smaller then the one of hand-reared cubs of the matching ages, but the difference was less pronounced than for the body mass (Fig. 2). The ratio of body mass vs. body length was greater in hand-reared compared to mother-nursed cubs. DISCUSSION The 1st feed formula was composed with no attempt to imitate the bear milk. It had high carbohydrate content and was relatively low in fat and proteins. This formula holds 50% of energy in carbohydrates and 39% in fats. The pneumonia of the cub 2 explains her lower consumption and mass gain. Hock (1966) reported on the growth of a three weeks old American black bear cub on a diet of evaporated milk and water in a 1:2 ratio. That cub consumed 125 ml food (42 g dry mass or 515 kJ) a day and gained 119 g in one day, so 4.33 kJ/g gain were needed. This conversion record is not fully comparable as the mass gain was measured in a single day where the last meal could have had significant influence. An attempt was made to make the 2nd feed formula more similar to bear milk by adding fat (cream) and proteins (yolk). The fats became the major energy source (66%) while carbohydrates dropped to 21%. The consumption grew 22%, and the mass gain 2.2 times. While the conversion of food mass to body mass was 37% better, the caloric conversion was slightly worse. Rationing of meals for cubs 3 and 4 decreased their mass gain as well as the conversion rate. Butterworth (1969) was feeding an American black bear cub for the first 8 weeks of its life with the formula consisting of equal parts of evaporated milk, water and light cream fortified with egg yolks. Until the same estimated age (i.e.. 82 days) Hulley (1976) was feeding American black bear cubs with evaporated milk diluted with water in ratio 1:3 for the first 2 days, 1:2 next 5 days, and 1:1 till the transfer to solid food. It is interesting to note that his cubs started to drink from the dish at the age of 72 days, i. e. almost the same as ours. The American black bear cub reported by Hock (1966) at the age of 80 days showed a mass gain of 137 g/day with 2001 kJ ingested. Thus 14.6 kJ/g gain were needed, i. e. 3.7 times more than at the age of 20 days, but 3.2 times less than our cubs at the comparable age. The 3rd feed formula was a simulation of brown bear milk as described by Jenness et al. (1977). With over 40% of solids the milk of bears has high caloric content comparable only with the milk of cetaceans, pinnipeds, and certain rodents, but not with those of other fissipeds. In the brown bear milk Jenness et al.(1977) found 22% of fats, 11% of proteins and only 0.6% lactose. In our mixture the fat content (from vegetable oil) was the major source of energy (82%) with no carbohydrate energy at all. Despite a slight decrease in daily consumption the mass gain with this food was the highest and conversion the best. Compared with the mean values for all used feed formulas (including this one) the gain was 1.76 times higher, the conversion per mass of food was 5.0 times, and per caloric value 2.8 times above the average. The practically double amounts of food consumed by cubs 1 and 2 showed negative conversion results apparently because of incomplete digestion. The mixture of simulated bear milk with full cow milk showed 1.8 times smaller day mass gain than the pure simulated bear milk. Also the conversion was 2.2 times worse considering the amount of food and 1.4 times worse considering the caloric value. The higher amounts of food consumed by cubs 1 and 2 showed not only the negative conversion results but also the smaller day mass gains. The cub 5 came to our hands in very poor condition in the age of 3 months with only 1/3 of average mass of other 4 cubs at that date (2730 vs 8500 g). However, with the below average consumption (729 ml) she had the highest day gains (138 g) with the best conversion i. e. 5.3 g food or 38.6 kJ per g mass gain. The 5th feed formula was again rich in starch (40% of energy) and since the age of 135 days was the first solid food; the suckling was abandoned in feeding practice. Hulley (1976) transferred his American black bear cubs to solid food at the age of 107 days. The daily gain and the conversion of caloric value didn't change but the conversion per mass of food was 1.58 times worse. By the end of this period (age 186 days) all cubs approached each other in their masss: i. e. the difference between the smallest cub 5 and the largest cub 1 was 27% (mean mass = 18800 g). During 1.5 month while free living and free feeding the cub 3 gained only 20% less mass than was the average with our food mixtures. The exact composition, volumes and caloric value of natural foods remained unknown. However, it seems that the social, rather than nutritional, force brought 3 cubs back to people and captivity. The 6th feed formula (the standard Zagreb-Zoo adult bear food) was the richest in carbohydrates and the poorest in fat of all used foods and showed the lowest conversion results. In this formula 76% of energy derives from starch and 25 from fat. However, over the period of 1165 days of it's use the rates of conversion were considerably changing. In the 1st period, i. e. till the age of 10 months the average daily mass gains were the highest recorded but the conversion of food mass was 2.7 times, and of caloric value 3.1 times worse than that of the feed formula 5. During the 2nd year of life on the entirely same food, the mass gain dropped 1.61 times (the same as conversion indices) compared to the first 84 days of feed formula 6. In the cubs' third and fourth year the gain was further 2.2 times smaller. This was due to the completion of the subadult phase of growing. Our data indicate four distinct periods considering the number of days necessary for doubling of body mass. Until the age of almost 4 months the mass doubled every 32 days, from the ages of 5 to 10 months every 135 days, in the second year of life every 375 days, and in the third and fourth year only after 1755 days. The influence of different feed formulas used was of relatively lesser importance to the mass doubling periods than the phase of growth itself. Rausch (1961) and Hock (1966) recorded that artificially fed American black bear cubs reached greater masses than the mother-nursed ones at the matching ages. From the measurements reported by these two authors and by Butterworth (1969) of another hand-reared American black bear cub (total 15 captive and 9 wild cubs >1 year age), we calculated the linear regression growth lines for both groups. In the start of artificial feeding (ages 0.5-3 months) orphaned cubs were, in average, smaller than mother nursed ones. We assume this is because of common malnutrition of orphaned cubs taken in emergency to artificial feeding. However, around the age of 105 days the masses of captive and wild American black bear cubs were the same, and later the hand-raised cubs were increasingly larger until they became double that of wild cubs around the age of 310 days. This is corroborated with our data on European brown bears (Table 2 and Fig. 1). However, when we analyzed the growth of body length the difference in favour of hand-reared vs. mother-nursed cubs was smaller (Fig. 2). This indicates that unrestricted feeding resulted in greater body mass as a combination of faster general growth (i. e. length) and of larger fat deposits. Oftedal and Gittleman (1989) demonstrated bears vs. other terrestrial carnivores to be on the extreme lower side considering the litter vs. mother mass, litter metabolic body size, litter postnatal growth rate vs. mother size, and milk energy output at peak lactation. They discuss three contributing factors: hibernation, herbivorism and maternalism. Though limitations imposed by hibernation influenced small litter mass and low milk energy output it is to some degree compensated by high milk energy content from maternal body fat while conserving maternal lean body mass. Herbivorous/omnivorous diet on foods of low digestible energy content may limit mothers ability to acquire and process sufficient energy to support high milk output and rapid postnatal litter growth. Single parental care (maternalism) vs. biparental or communal caring systems (e.g., candies) may increase the amount of energy in acquiring prey or providing protection for the young. According to data presented by Oftedal and Gittleman (1989) American black bear cub nursed by his mother consumes at the peak of lactation, in the age of 75 days, 168 g of milk per day, or 5.8% of body mass. Our hand-raised brown bear cubs at the same age consumed an average of 707 g/day of feed formula, or 11.1% of their mean body mass. As stated earlier they were also, most of the time, more than twice larger than wild brown bear cubs. The similar was documented for American black bears (Rausch, 1961). We speculate that bears resumed evolutionary genetic potential for utilization of larger amounts and higher quality of food and to convert it to body tissues. The striking faster growth rate with artificial feeding may be a part of general phenotypic reactivity exhibited by bears worldwide as a response to different environmental conditions. It also gives further evidence of an incomplete evolutionary transition from carnivore to herbivore diet. CONCLUSIONS 1. The high content of fat in the food of nursing brown bear cubs showed as good, and high content of carbohydrates as bad indicator. 2. Until the age of 4 months the cubs doubled their mass every 32 days, from the ages of 5 to 10 months every 135 days, in the second year of life every 375 days, and in the third and fourth year only after 1755 days. The mass doubling periods were more influenced by the phase of growth than by the feed formulas used. 3. Five hand-reared cubs were 1.3-2.7 times larger than 17 measured wild cubs at matching ages. In the unrestricted feeding they also consumed roughly double (1.9) amount of food than the mother would be able to provide them. 4. European brown bears seem to be able to eat larger amounts of more adequate food and consequently to grow larger. Limitations imposed by the birth during the mother's hibernation and by mainly herbivorous diet later in life influence their growth rate and the final body size. The environmental pressure has marked influence on the expression of genetic potential in bears, in which they exhibit the great phenotypic adaptability to live with the given opportunities. ACKNOWLEDGMENTS We are grateful to the Delnice, Vrbovsko, and Ogulin Forest Enterprises, Risnjak National Park, and Zagreb Zoological Garden for the help in obtaining the bear cubs and their maintenance. We gratefully acknowledge the enthusiastic help provided by A. Frkovi} and the students of the Veterinary Faculty in Zagreb, particularly V. Dabanovi}. We also thank R. Flowers for reviewing an earlier draft of this paper. REFERENCES Butterworth, B. B. Postnatal growth and development of Ursus americanus. JOURNAL OF MAMMALOGY, 50:615-616, 1969. Frkovi}, A. Tragom markiranih medvjedi}a iz Lividrage u Gorskom kotaru. PRIRODA, 71:72-73, 1983. Hock, R. J. Growth rate of newborn black bear cubs(Ursus americanus). GROWTH, 30:339-347, 1966. Hulley, J. T. Hand-rearing American black bear cubs Ursus americanus at Toronto Zoo. INTERNATIONAL ZOO YEARBOOK, 16:202-205, 1976. Jenness, R.; Erickson, A. W.; Craighead, J. J.. Some comparative aspects of milk from four species of bears. JOURNAL OF MAMMALOGY, 53:34-47, 1972. Oftedal, O. T.; Gittleman, J. L. Patterns of energy output during reproduction in carnivores. Pp. 355-378, in CARNIVORE BEHAVIOR, ECOLOGY, AND EVOLUTION. J. L. Gittleman, ed. Chapman and Hall, London, 620 pp., 1989. Rausch, R. L. Notes on the black bear, Ursus americanus Pallas, in Alaska, with particular reference to dentition and growth. ZEITSCHRIFT FUR SAUGETIRKUNDE, 26:77-107, 1961. 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