F:\ALCES\Supp2\PAGEMA~1\Rus 16s ALCES SUPPL. 2, 2002 KUZNETSOV - FAECES AS INDICATORS OF MOOSE ACTIVITY 71 FAECES AS INDICATORS OF MOOSE ACTIVITY AND ROLE IN ECOSYSTEMS German V. Kuznetsov Institute of Evolutionary Animal Morphology and Ecology, Russian Academy of Science, 117071, Moscow, Russia ABSTRACT: Faeces can serve as a significant index of spatial distribution of moose and provide estimates of moose population composition. Using faeces of moose to gather data for determining forage pressure and level of removal of plants, territorial distribution, population density, traces of activity, and other population indices can be regarded as a simple and reliable method for investigation of moose ecology. ALCES SUPPLEMENT 2: 71-76 (2002) Key words: ecosystems, faeces, forage digestibility, moose, plant mass, Valdai By consuming plants, excreting faeces and urine into the environment, maintaining metabolic processes of the body, and through diurnal and seasonal displacements, moose are involved in transfer of energy and me- tabolism of matter in forest ecosystems (Kusnetsov 1976). Faeces can serve as a significant index of spatial distribution (Yurgenson 1961, 1970), the amount of consumed production (Semenov-Tyan- Shansky 1948, Kuznetsov 1975), transfer of energy (Kuznetsov 1976), and other forms of moose involvement in ecosystems. Estimation of ungulate faeces as a method for the evaluation of the numbers of moose and spatial distribution first origi- nated in the USA as early as the 1950s, and in the USSR this method was first applied in 1957 by P.B.Yurgenson, who proposed an original variant based on the common prin- ciples of the MacQuane method adopting an almost constant number of defaecations per day in ungulates (Yurgenson 1970). Subsequently, faeces as an indicator were successfully used to determine the foraging pressure of moose and their winter distribu- tion in various hunting grounds and habitats of European Russia (Pivovarova 1965, Ivanova 1967). Our studies in Valdai based on an annual estimate of moose faeces along a 3 km transect with constant sam- pling sites (300 m2 every 50 metres), dem- onstrated varying pressure of moose on particular habitats. In fact, the amount of faeces, on average, on the sample site over the year reached 147 ± 46 pellets in the pine forest with cowberry, 85 ± 27 pellets in pine forest with green moss, 72 ± 11 in the pine forest with sphagnum, 60 ± 10 in spruce forest with herbs, 59 ± 9 in a mixed sphag- num forest, 49 ± 11 in a spruce forest with green moss, and 5 ± 1.5 in a mixed sphag- num forest (Fig. 1). Over the 20 years of observations (1970–1990), the greatest number of faeces in the pine forest with sphagnum was recorded > 7 times, in the pine forest with cowberry 5 times, in the pine forest with green moss 2 times, and in the spruce forest and other types of forest 1 time, which is indicative of different trophic and topical pressure of moose in these types of forest. Interestingly, in Valdai we re- corded dissimilar rates of visitation by moose of particular habitats (Fig. 2); this is true of other regions as well (the Tula Region). Thus, depending on the amount of forage FAECES AS INDICATORS OF MOOSE ACTIVITY - KUZNETSOV ALCES SUPPL. 2, 2002 74 Pivovarova 1965). Such data make it pos- sible to estimate the involvement of moose in transfer of energy in different biocenoses. It is important to take into account the existence on the same ranges of the migrat- ing and sedentary part of moose populations (Knorre 1959, Baskin 1984), which influ- ences the complex biological nature of the role of moose in the ecosystems. The method for determining the amount of plant mass consumed by moose accord- ing to their faeces was first applied by Semenov-Tyan-Shansky (1948). The use of this method in nature necessitates data on the amount of faeces (in dry weight) excreted by moose in a particular area, the period of time over which the moose defaecated, and the coefficient of forage digestibility. On the whole, it can be as- sumed that the mean annual coefficient of food digestibility in moose is close to 70%. It does not exceed 50–55% in winter and reaches up to 84% in summer (Ivanova and Simakov 1975; Kuznetsov 1975, 1976). Thus, on obtaining data on the amount of faeces excreted by moose (dry weight) over the year and in a definite area (the constant sample sites are preliminarily cleared of old faeces) and knowing the coefficient of forage digestibility, one can easily estimate the amount of forage con- sumed by moose over the year in a particu- lar territory, using the formula for the di- gestibility coefficient (Kuznetsov 1975, 1976). Subsequently, Abaturov (1980, 1984) proposed estimating the amount of vegeta- tion consumed according to dry weight of the accumulated faeces and coefficient of forage digestibility by the formula: C = (Fx100)/(100-B); where, C = consumed phytomass (dry weight), F = dry weight of the faeces, and B = coefficient of digestibil- ity of the phytomass of the animal species (%); i.e., a mathematically proposed form is deduced for the digestibility coefficient. In this respect, the method of Milner (1967) is also of interest. Milner estimated the amount of forage consumed by animals from the ratio of the weight of faeces to the forage consumed as established in the labo- ratory. Subsequently, by estimating the amount of faeces left by animals over a certain period, one can determine the amount of the forage consumed: C = F/(F/C) (Dinesman and Khodashova 1974). In the field, the amount of forage con- sumed by moose was determined from their faeces in the population of the Byelorussian Lake Region (Dunin 1989) and Valdai (Kuznetsov 1975, 1976). This method makes it possible to objectively evaluate the re- moval of forage by moose in a definite territory without estimation of their popula- tion. Along with that, a number of authors (Zlotin and Khodashova 1974, Abaturov 1984) indicate that the accuracy of the results can be affected by the activity of microorganisms and coprophagic animals, and also the effect of the physical factors of the environment in particular geographical zones. For instance, in the subarctic, con- ditions for the rate of decomposition of moose faeces is low. The mass of faeces over 4 years declines by 53% (Malafeev and Kryazhimsky 1990). According to our data, under Valdai conditions, the mass of moose faeces decreases by 64% over 4 years, and by 89% over 6 years. One can estimate the population density of moose according to the amount of faeces deposited by a moose population over a certain time in a particular territory (Yurgenson 1961, 1970; Kuznetsov 1976). Based on the fact that every adult moose over a year excretes 2 kg of faeces (dry weight) every day (this is in conformity with the data of Knorre and Knorre 1959), we conclude that the population density of moose in Valdai, the amount of faeces excreted by moose in this area over 1 year being 2.9 kg/ ha, represents 1 individual per 252 ha; i.e., 4 individuals per 1,000 ha (Kuznetsov 1976). FAECES AS INDICATORS OF MOOSE ACTIVITY - KUZNETSOV ALCES SUPPL. 2, 2002 76 Pages 281–287 in A. G. Bannikov, edi- tor. Biology and harvest of moose. Rosselkhozizdat, Moscow, Russia. (In Russian). , and A. F. SIMAKOV. 1975. Protein metabolism. Pages 63–127 in Com- parative physiological study of protein and mineral metabolism in ruminants under northern conditions. VINITI, Moscow, Russia. (In Russian). KNORRE, E. P. 1959. Ecology of moose. Proceedings of the Pechoro-Ilych State Reserve 7:5–22. (In Russian). , and E. K. KNORRE. 1959. Inves- tigations of some physiological charac- teristics of moose. 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