* Chapter 13:
General Features of Diet.š Influence on Prey Populations. Competitors and Scavengers.
An adult tiger "over a short period of time...is capable of consuming a large amount of meat.š For example, once a large male, without changing his position (lying on his stomach), ate a roe deer that he had knocked down without leaving any part of it uneaten...Another male ate 75% of the carcass of a "seven-point" Manchurian deer (not less than 80 kg of meat and bones) over a period of less than 3 days " (Fig. 40) (Yudakov 1973, p. 93).š An estimate of the normal amount of food required by tigers is made more difficult by the fact that a prey animal is caught by the tiger within a time period that is neither preset nor equal to a regular time unit.š It is already possible to make a judgment about this simply by considering the fluctuations in the success rate of the tigers' hunts, which have been examined in the previous chapter.š For the tiger "it is natural to go hungry for up to 10 or more days.š The time between successive prey kills ranges from several hours up to 11 days, averaging 5 days" (Yudakov 1974, p. 355).
In order to present approximate quantitative indices characterizing the tiger's diet, it is advisable first to utilize data from those tracking episodes where the segment of the series of footprints had a sufficiently clear temporal referencing.š By summing together such segments for male tigers, we will produce a characterization of their lives over a 41-day period (460 km of tracking).š Over that interval of time, there were ten prey kills with one animal being an adult female wild boar that had been killed by a tigress but that was eaten by her together with a male tiger during their joint travels.š There were eight wild boar (7 of which were yearlings) and two Manchurian deer (an adult bull and an adult female) among the ten prey kills.š With the exception of one of the Manchurian deer, more than 75% of the weight of each animal was consumed.š The total quantity of meat and bones consumed was approximately 300 kg.š If we extrapolate this result to an entire year, we arrive at a value of 2,600-2,700 kg.š This means that, on average, a male tiger eats only somewhat more than 50 kg of meat per week.
A calculation based on the total aggregate of the data is still more approximate in nature.š It is based on the use of an index of the average daily travel distance, with the aid of which it is possible to roughly estimate the total number of 24-hour periods in the life of tigers that have been characterized by our tracking.š Such a computation for male tigers gave results that are extremely close to those that we have just examined.
There were 11 prey kills for 28 24-hour periods in the life of tigresses (219 km of tracking).š The species composition of their prey is more diverse than in the previous case: six wild boar (two adults, four piglets); one yearling Manchurian deer (saek); one musk deer, and one dog. Such, we may assume, is the monthly ration for one individual female.š It amounts to approximately 250 kg of meat.š This index exceeds that for males, however, a calculation based on the total aggregate of data leads to the opposite conclusion: the female's demand for food is significantly less than that of males.š In any event, estimates that were arrived at using a variety of methods are similar and, apparently, reflect the true orders of magnitude.š If over the course of a week, a tiger requires more than 60 kg of meat, then for an entire year this comes to slightly less than 3,000 kg.
Figure 40.š The remains of a Manchurian deer killed by the male "Emperor" tiger.
Our estimates are very close to analogous indices calculated by L. G. Kaplanov (1948).š According to the data of this investigator, an individual tiger consumes approximately 3,000 kg of meat over the course of a year, which corresponds to 30 nominal prey animals each weighing 100 kg.š Since young animals predominate among the tigers' prey, the actual number of prey turns out to be considerably greater than these 30 nominal animals: in our case, it is 90 individuals for males, while for females, it is greater than 120 individuals.š Here an estimate made using the total aggregate of the data also partially equalizes the difference between males and females, with the number of individuals captured per year becoming less and approximating the estimate that has been published earlier by A. G. Yudakov (1973): 70-75 animals.š The difference between males and females reflects not so much the total amount of food eaten by them, so much as the dimensions of the animals that comprise the prey.š "An adult female kills a greater number of animals, but smaller animals predominate among her prey" (Yudakov 1973, p. 93).
During the course of all the years of our study, the main prey of tigers in the study region was wild boar (Table 18).š Not less than half of the prey, on average about 60%, was attributed to this species in each of the seasons.š Manchurian deer constituted only a quarter of all the animals killed by tigers.š Other species occupied a secondary position in the tiger's diet.š In comparing the data on the population sizes of ungulates in the long-term study site (cf., Table 4) with the relative importance of different prey species in the diet of the tiger, it is clear that these predators kill wild boar more often than would be expected based on the ratios of the densities of prey populations.š Thus, in the winter of 1970-1971, Manchurian deer were even more numerous than wild boar in the study region, but half of the animals captured by tigers were attributed to the latter species, while Manchurian deer made up less than a quarter of all prey killed.š These differences find an explanation in the difference in the success rates of these predators' hunts on Manchurian deer and wild boar, a fact that has already been mentioned.
Prey composition of tigers in the long-term study area (based on finds of prey remains supplemented data derived from an examination of the tigers' excrement).
š
š
Winter Season
Total Number of Prey
Wild Boar
š Manchurian Deer
šš Roe Deer
š Musk Deer
š Himalayan Bear
š Dog
š Total
% Total
% Total
% Total
% Total
% Total
% 1970/71 14 7(2) 50 3 21.4 - - 1 7.1 2 14.3 1 7.1 1971/72 22 13(4) 59.1 5(1) 22.7 3 13.6 - - - - 1 4.6 1972/73 28 18(2) 64.3 8 28.5 1 3.6 1 3.6 - - - - Total 64(9) 38(8) 59.4 16(1) 25.0 4 6.3 2 3.1 2 3.1 2 3.1 Note:š The number of data points included in the total data set based on the analysis of tigers' excrement is given in parentheses.š The latter source of data was taken into account only in those instances were these data gave information on prey that we did not discover during our tracking.
Selectivity is also manifested in the ratio of the age composition, and also partly of the sex ratio of the prey.š The percentage of yearlings among the wild boar killed by tigers is considerably higher than its percentage in the population (Table 19).š The same could also be justifiably said of the "sayok" (or yearling) Manchurian deer (Table 20).š The proportion of adult bulls of the Manchurian deer that are part of the prey composition of tigers is relatively low.š Apparently, this sex and age group suffers the least from the tiger.š The effect of the predator is directed in the present instance, as can be seen from these data, above all toward the utilization of the annual population increase.š With the transition of young ungulates to older age groups, their risk of death from the tiger decreases substantially.
On the basis of the available data we can only make approximate calculations with respect to a general estimation of the annual losses to the wild boar and Manchurian deer populations from the tiger.š A sufficiently reliable estimate can be presented only for the snow season (five months--from the middle of November to the middle of April).š The proportion of individuals removed from the populations of ungulates by the tiger is calculated for an area of approximately 900 km2, where we carried out continuous observations.š A male tiger and a female, which had two cubs during the first half of this time period and one cub in the second half, lived here over the course of three winter seasons.š In the final season of our observations another male transited through this territory, and, toward spring, we noted the footprints of yet another female, which had two cubs.š Provisionally, it is possible to accept an estimate of three adult individuals as inhabiting the area in question.
Age composition of the wild boar population (based on encounters with these animals) and the percentages of animals of different ages that became the prey of tigers.
š
Composition of the Sample
Number of Individuals Counted
Adults
Yearlings
Total
% Total
% Live Wild Boar Observed
144 53 36.8 91 63.2 Prey of the Tiger
30 7 23.3 23 76.7 Age composition of the Manchurian deer population (based on encounters with these animals) and the percentages of animals of different ages that became the prey of tigers.
š
šš
Composition of the Sample Number of Individuals Counted Adult Males
Adult Females
Yearlings Total % Total % Total % Live Manchurian Deer Observed
93 22 23.7 28 30.1 43 46.2 Prey of the Tiger
15 2 13.3 4 26.7 9 60.0
The number of 24-hour periods in the life of the tigers that we tracked is determined by the total tracking length and the average 24-hour run.š Estimates made using single segments of the tigers' journeys are, in our opinion, less informative here due to the relatively small amount of data available in the present case, and also due to fluctuations in the success of the hunts in winters with varying snow conditions.š Summing up for 148 "estimated" 24- hour periods in the lives of all the animals that we had under observation, they had killed 19 wild boar and 7 Manchurian deer.š These data may conditionally be ascribed to a single tiger's life.š Taking into consideration the differences in prey composition and in the quantity of prey taken by male and female tigers, it is possible to conclude that the three tigers captured up to 60 wild boar and 20 Manchurian deer over the course of five winter months.
The mean indices of the population densities of wild boar and Manchurian deer in the long-term study site over the course of three winter seasons were 4.2-5.0 and 3.5-4.0 individuals per 10 km2, respectively.š Comparing these figures with the results produced above, it is possible to conclude that over the course of a single snow season, the tigers in the sector under our observation removed on average up to 14% of the wild boar population and up to 6% of the Manchurian deer population.š The annual losses to the populations of prey are approximately twice as large.š With regard to wild boar, the magnitudes of these estimates that we have calculated are very close to the estimates which A. G. Yudakov (1973) made on the basis of preliminary computations from our data, but the percentage removed from the Manchurian deer population is significantly less that the proportion originally accepted (i.e., in Yudakov's earlier estimates).
Domestic animals in that part of Sikhote-Alin where we conducted our observations are an insignificant element in the diet of tigers.š According to the information that we succeeded in gathering, the number of cases of attacks on livestock and dogs over the course of more than half a century amounts to several 10s of events (23 instances were recorded, Table 21).š Moreover, during the years of our work, only dogs perished due to tigers.š Almost all these cases took place in the taiga when dogs were used during hunting.
Only the brown bear can play the role of a competitor with the tiger in our study region, and then mainly only during a very brief period of the year--early spring.š After their emergence from their dens, which are concentrated near the divides of watersheds, the bears descend into the lower zones of the mountains, where they begin hunting wild boar and Manchurian deer.š We had already noted their first appearance in the middle of March in areas where the tigers constantly traveled.š It was precisely here that we observed (by tracking) a hunt by a brown bear on Manchurian deer at the end of this month.š We twice found the remains of ungulates that had been killed by brown bears during the spring season: in March, the remains of a musk deer and in April, the remains of an adult Manchurian deer.š In April, we observed a case where a Himalayan bear (we observed him visually) was feeding on the remains of a Manchurian deer that had been killed by a tiger.š In autumn, the footprints of bears were common up until the beginning of December.š Our latest (for the season) encounter with the footprints of a brown bear took place on December 21.š In the pre-winter period, only one animal that had fallen prey to a brown bear was discovered: this was a wild boar killed at the end of November.
Incidents of tiger predation on domesticated animals in the basin of the Malinovka River (up to 1973).
š šš
Date
Type of Animal Killed
Number of Individuals
Location and Circumstances of the Attack
Author of the Report
1912 š
š
Horse
1 š
š
Village of Savinovka (in a stable), owner: N. Romanyuk
A.M. Urbanovskii
š
š
1920 š
š
Bull
š
š
1 š
š
Near the settlement of Mezhdurech'e, owner: Vigovskii
V.A. Yarich
š
š
1938 Dog
1 Near the settlement of Martynova Polyana, owner: Vorozhbit, the tiger killed the dog in his presence
N.M. Vorozhbit
1951, summer Horse
3 Near the settlement of Polyana, in a pasture, one of them was a foal
N.A. Pervukhin
1951, November Dog 1 Near the settlement of Polyana, killed by a tiger in the presence of a hunter
N.A. Pervukhin
1952, February Cattle
3 Settlement of Polyana, in a stockyard
N.A. Pervukhin
1958 Dog 3 Upper reaches of the Malinovka River, during a hunt, owner: Drachuk
P.E. Asyrkin
1958 Cow
1 At a distance of 4 km from the settlement of Ariadnoe, in a pasture
A.M. Ivashchenko
1958 Horse
1 Near the settlement of Pozhiga
K.G. Abramov, 1965
Unknown
Dog 2 Pyatyi Stream (Bystraya River), owner: A. Prokopchuk
D.M. Dobroshevskii
Beginning of the 1960s, summer
Cow
1 Near the settlement of Pozhiga, owner: K.I. Darenskii
P.E. Asyrkin
1967 Dog 1 Bogdanov Stream (Gornaya River), killed in the presence of hunters by a tiger
A.V. Deryagin
1968, autumn
Dog 2 Orekhovka River, on a hunt
N.A. Pervukhin
1969, May Horse
1 Village of Lyubitovka
A.F. Novitskii
1970, autumn
Dog 2 Lazarev Stream (Titovka River), at an apiary, owner: Shchegolyuk
G.F. Gorokhov
1970, September Cow
1 Settlement of Pozhiga, in a pasture, owner: P.A.Toporkin
P.E. Asyrkin
Table 21 (cont.)
šš
Date Type of Animal Killed Number of Individuals Location and Circumstances of the Attack Author of the Report 1970, November
Dog 1 Upper reaches of the Malinovka River, on a hunt, owner: Artemenko
G.F.Gorokhov
1970, December Dog 1 Settlement of Martynova Polyana, owner:Volokotyuk
A.V. Deryagin
March 9, 1971 Dog 1 Pravyi Igristyi Stream (Gornaya River), found in the taiga without its owner
Tracking Data
November 15, 1971 Dog 1 Kammenyi Stream (Gornaya River), on a hunt, owner: A. Kiryushkin
Tracking Data
1971 Dog 1 Upper reaches of the Malinovka River, on a hunt, owner: G. I. Korzh
P.E. Asyrkin
1972, January Dog 2 Klinovka Stream (Orekhovka River), on a hunt, owner: Grigor'yev
T.L. Trofanchuk
1973, October Dog 1 Bystryi Stream (Orekhovka River), on a hunt
Information collected by A.G. Yudakov
In one case, we observed a direct encounter between a tiger and a lynx by monitoring their tracks.š The tigress, having encountered a lynx near a roe deer that the lynx had killed along her route, stopped, sat down, and then rushed to the site by leaping and drove the lynx away from her prey.
For our long-term study site, we found that the interactions between the tiger and scavengers-commensals that were examined in greatest detail by E. N. Matyushkin (1974) for the eastern macroslope of the Central Sikhote-Alin Mountains were exactly the same as those described by Matyushkin.š Here, the main beneficiaries of the remains of the tigers' prey were also the large-billed crow (Corvus macrorhynchus) and the raven (Corvus corax).š Among the other birds, that we observed at the prey killed by tigers were the jay (Garrulus glandarius), the Eastern blue magpie (Cyanopica cyana), the nuthatch (Sitta sp.), and the Siberian tit (Parus cinctus) (Fig. 41).š Without question, the Siberian jay (Perisoreus infaustus) also belongs among the scavengers of tigers' kills, although we did not observe it in such a situation on even as single occasion.š In places where this bird is common, it often falls into hunters' traps, to which it flies, being attracted by the bait.š Among mammalian scavengers, the Siberian weasel (Mustela sibirica) feeds more frequently than others on the tigers' kills.š We encountered the following scavengers on a single occasion each: a mink (Mustela lutreola)š (which was frightened away from the remains of a roe deer), a group of yellow-throated marten (Martes flavigula), a sable (Martes zibellina), a raccoon dog (Nyctereutes procyonoides) and a Himalayan bear.š In four cases, wild boar ate carrion (three times, it was the remains of a Manchurian deer and once the remains of a roe deer).š The remains of the wild boar themselves, around which we once found their tracks, were not touched by the wild boar.š Wild boar ate not only carrion that was lying out in the open, but they also dug up carrion from under a thick layer of snow (Fig. 42).
An examination of the remains of 42 of the tiger's prey allows us to conclude that the plunderers of the tiger's prey misappropriated approximately 15% of that part of the prey that the main scavengers are capable of utilizing.
Figure 41.š A jay (Garrulus glandarius) feeding on the remains of a tiger's prey: a yearling wild boar.
Figure 42.š The remains of a roe deer killed by a tiger after having been dug out from under the snow by wild boar.
š
š
Copyright ¿ A. G. Yudakov,I. G. Nikolaev
Copyright ¿ K. Lofdahl, A. Shevlakov, 2004 (English translation)