* Chapter 12

Hunting Behavior and Success of the Tigers' Hunts

Different individual tigers can utilize one and the same prey carcass, but these predators always or almost always hunt alone.š We did not observe even a single instance of group hunting by adult animals during the entire period of our study (which also included the mating period).š Approximately two thirds of the hunts that we tracked took place on mountain slopes with the rest occurring in the valleys of rivers and streams.š Of the 40 tiger hunts that can be attributed with sufficient precision to a definite time interval, 25 (62.5%) took place during the dark portion of the 24-hour period.š In addition, it was precisely established in the case of 17 of these hunts that the hunt took place either in the pre-dawn or evening hours. We shall further separatelyšexamine cases of the hunt where prey were wild boar and Manchurian deer.

Tigers encountered their potential prey and made attempts to pursue them during their habitual travels throughout their territories.š Special "hunting" sectors along their routes could not be clearly distinguished.š Hunting sites on the routes of both male tigers and tigresses that lacked offspring were distributed fairly uniformly. The relative concentrations of these hunting sites, which are visible on the map (Fig. 28), are associated with the core of the family territory or with sites of long-term residency by a female with her cubs or by adult animals during the mating period.š In the presence of a sufficient density of prey species and of their relatively uniform distribution, which is characteristic of the study region, this way of exploiting their territory during hunts may be considered typical of the tigers.š Favorite hunting sites are apparent on the territories of these predators only under conditions of distinctly expressed seasonal concentrations of ungulates in definite isolated places in the landscape, as has been observed for the Lazovskii State Nature Reserve by V. I. Zhivotchenko (1976).

The frequency of hunting attempts along the tigers' routes is very variable.š The intervals between such attempts range from several hundreds of meters to several tens of kilometers.š In the case of males, one hunt occurs, on average, for every 18 km of the tiger's path, while for females, one hunt takes place for every 12 km of the tigress' route.š Successful hunts during our tracking were recorded approximately 2.5 times less frequently (for 47 km of the tigers' travels in the case of males and for 27 km of the tigress' journeys).š Encounters between predators and potential prey are much more frequent than hunting attempts.š Thus, on March 7, 1972, a tigress frightened off wild boar three times (in one case, a small herd and in two instances, single animals), roe deer twice (two animals in each instance), and Manchurian deer once along a route of length 12 km walked by the tigress in the basin of the Orekhovka River during daytime.š There were no attempts to attack the animals in these instances, since the animals detected the predator in good time at a distance that precluded a successful pursuit.

Figure 28. The distribution of tiger hunts on wild boar and Manchurian deer in the winter seasons of 1971-1973.

Hunts on wild boar: 1--successful; 2--unsuccessful.š Hunts on Manchurian deer: 3.--successful; 4--unsuccessful (the diagram does not reflect the mean values of the success rates of hunts, since here we indicate also sites where we found old carcasses not having any relation to the period in the life of the animals that was characterized by tracking)

The stimulus for the initiation of a hunt is usually the nearness of the prey animal itself rather than the tiger's discovery of fresh tracks made by the prey.š In only 7 of 39 observed cases of tigers' hunts on wild boar did the tigers begin their pursuit of prey when they were taking their cue, in all probability, only from the fresh tracks of the animal.š With regard to Manchurian deer, similar reactions were noted to be still more rare: 2 instances out of 40 hunts.

It is generally accepted that tigers, like other felids, mainly discover their prey with the aid of sound and sight.š However, it is practically impossible to estimate the maximum distance at which tigers can hear, still less see, a possible prey animal by using the tigers' tracks.š With respect to the effect of olfaction in such situations, we present below several specific observations, when the circumstances excluded the discovery of objects that interested the tigers by using sound or sight.š Thus, a female, walking along a road during the dark part of the 24-hr period, abruptly turned off the road and, walking over virgin snow for a distance of 130 m, approached the freshly-removed skin of a Manchurian deer, which had been prepared by hunters.š In another case, a male tiger, after making a sharp turn away from his route and after going a distance of 130 m, approached three wild boar dens, which had been abandoned by the animals shortly before that time.š In both cases, the tigers returned to their former routes by following their own old tracks.š A similar reaction of a predator to a wild boar that was present in a den was observed once at a distance of 195 m from the den.š The maximum distance at which the tigers discovered Manchurian deer and roe deer in the ungulates' laying places turned out to be still greater: 200 m and 230 m, respectively.š However, here too there was reason to believe that they were guided by olfaction and not by sound.

Animals that are the possible prey of tigers react in different ways to footprints of these predators and to the scent of the tigers.š Manchurian deer, for example, do not manifest any indications of unease when they run across fresh tiger tracks.š Wild boar usually behave in a similar fashion.š We observed on one occasion, however, that wild boar, after approaching their den for the night and having discovered that a tiger had lain there not long before their arrival, abandoned this place.š In one case, roe deer immediately began leaping away upon crossing fresh tiger tracks.š Various reactions were observed in such situations in the case of bears.š One brown bear, which was staying in an area permanently inhabited by tigers, clearly felt himself to be the complete master in that place.š Another brown bear, once abruptly turned away from his former path upon his encounter with tiger tracks.š But a large, apparently male, Himalayan (or Asiatic black) bear (which we observed visually), like the brown bear that has already been mentioned, clearly did not fear the presence of tigers.š He walked along the tiger's tracks and rested in the same wild boar den as did the tiger.š Thus, the tigers' fresh footprints do not produce a strong sense of unease in the tiger's potential prey in the majority of cases, a fact which is very important for ensuring regularly successful hunts by these predators in areas where tigers live on a permanent basis.

Observations of hunts by tigers on wild boar and Manchurian deer do not provide a basis for speaking of a distinct preference by tigers for one type of prey over another.š In conditions that are favorable for attack, charges by these predators after one or the other species are, apparently, equally probable.š However, these hunts take place in somewhat different ways, and it is therefore more convenient to examine them separately.

Hunts on Wild Boar:


Tigers pursue wild boar mainly on mountain slopes, which are inhabited by these ungulates primarily during years when there are good harvests of their main foods.š Out of 39 hunts on wild boar, 34 (87.2%) took place on slopes or on the foothills of slopes.š Tigers encounter wild boar in valley habitats most often during periods in which the wild boar are concentrated in areas where horsetails (Equisetum spp.) grow abundantly.š Tigers often use wild boar trails when making their journeys, as a consequence of which the probability of encounters with these prey increases.š It is under precisely such conditions that the predators discover the wild boar in approximately half of the cases of hunting that we investigated.

The attacks took place both in darkness and in daylight (Table 15).š We succeeded in establishing in eight instances that the hunts probably occurred during daylight, however two of these cases took place during the morning hours and two occurred during the evening.š We observed seven nocturnal hunts, with three of these occurring soon after the arrival of darkness and one occurring before dawn.š Direct observations on wild boar in the long-term study site revealed that during the daytime these animals, as a rule, forage or travel around searching for food, but with the arrival of dusk, they lie down in a den, where they rest until 9:00-11:00 hours on the following day (Fig. 29).š Tigers either attacked foraging animals or attacked them in the wild boars' laying places (i.e., in their den), thus matching their behavior to the 24-hour activity rhythm of the wild boar.š Instances of hunts on foraging wild boar were observed significantly more often (Table 17), which allows us to conclude that a somewhat higher number of daytime hunts (versus nighttime hunts) represents the normal pattern.

Table 15.
Frequencies of daytime and nighttime hunts by tigers and the nature of prey activity at the moment of the predator's attack.

Object of the Tiger's Hunt Total Number of Hunts Time During Which Hunt Took Place Activity of the Prey
Daytime Night Total Number of Animals Subjected to Attack Feeding Lying Down
Abs. % Abs. % Abs. % Abs. %
Wild Boar 15 8 53.3 7 46.7 34 20 58.8 14 41.2
Manchurian Deer 20 4 20 16 80 34 22 64.7 12 25.3

The minimum, maximum, and mean distances at which these predators discovered wild boar (for 11 hunts) were: 65 m, 290 m and 194 m, respectively.š Despite the great diversity of situations making up each individual case, the tigers' methods of stalking prey generally followed a stereotyped pattern.š This pattern included either a hunt "from the tiger's path", when up until the tiger's sudden spring nothing in the characteristics of the tiger's tracks indicates an attack in the immediate future, or the stalk itself--with laying places, stops, the alternation of ordinary strides with slow, short steps with these sometimes changing to a fast trot before the final spring toward the prey.š During hunts on wild boar, attacks "from the tiger's path" were observed in 16 instances out of 39 hunts (41%).š Hunts involving the use of laying places, stops, and slow approaches to the prey were tracked by us on 13 occasions (33.3%).š Sometimes tigers undertake flanking maneuvers before the attack, the goal of which is, apparently, an approach to the object of the hunt from the downwind side or a search for a more convenient position for a spring.š While hunting on wild boar, the predators undertook such flanking maneuvers in 8 cases (20.5%). The use of an ambush by the tigers was observed only three times; one time, the tiger awaited the approach of a foraging wild boar, having concealed itself behind a Korean pine tree. On two occasions, it was established from the tigers' tracks that the predators had hidden themselves in "dens" (made by wild boar).š Twice during their stalking of wild boar, the tigers crossed over short (up to 8 m) distances by crawling, leaving a characteristic trench behind them in the snow.š In accordance with the circumstances of a particular hunt, we also happened to observe the successive use of different methods of stalking.

The distances from which the predators succeeded in approaching wild boar before completing their final charge varied: from 4 m up to 60 m (Table 16); the mean value (for 21 hunts) was 20.3 m.š The length of the tracks resulting from the tiger chasing after its prey (i.e., the leaps) fluctuated from 10 m up to 560 m (Fig. 30).š In only one case did the predator charge after wild boar over a distance of almost 900 m.š If we do not take into account this sharply deviating value, the mean index for the length of the chase for 29 hunts (both successful and unsuccessful) is 137 m.š Without the inclusion of three cases where pursuit occurred over a distance of approximately half a kilometer, which are also exceptional, the mean index decreases to 84 m.

Figure 29. A "den" made by wild boar located along the south-facing slope of the right bank of Khvoinyi Stream.

Table 16.
Distances between tigers and their prey before the tiger's charge and the distances over which tigers pursued their prey for both successful and unsuccessful hunts

Object of the Hunt Result of the Hunt Age of the Prey Number of Hunts Distance Between Tiger and Prey Before the Tiger's Charge (m) Number of Hunts Length of the Hunt (m)
Min. Max. Mean Min. Max. Mean
Wild Boar Successful Adults 5 8 15 10 5 30 135(900)* 67
Yearlings 8 4 30 16 13 10 150(390) 53
Unsuccessful Adults & Yearlings 8 4 60 31 11 15 560 200
ฺ้ภยาษ Successful Adults 4 3 22 13 4 48 74 59
Yearlings 5 3 18 11 6 19 40 31
Unsuccessful Adults & Yearlings 15 10 80 34 16 10 260 72
*In parentheses, we present sharply deviating maximum values that were observed only once.š The respective mean indices are calculated without taking these values into account.

The positions of the attacking predator and the prey in relation to the topography were variable.š The tigers were located above the prey in 15 instances (48.4%), and they were situated below the prey in 5 cases (16.1%).š Tigers attacked wild boar 11 times (35.5%) when the tigers were at approximately the same elevation as their prey.

Piglets and gilts (piglets that are 4-9 months old) that were overtaken by tigers were killed on the spot.š When adult wild boar was the prey, the wild boar dragged the predator along behind them in two out of five cases.š Once, the struggle of a tiger with a large female wild boar took place along a steep slope with the animals traveling 15 m downward along the slope; the wild boar was killed near a large dead standing tree that obstructed the prey's path.š In another instance, the short, overlapping leaps made by a tiger and a female wild boar, which later changed to the intermixed tracks made during their struggle, extended downward along the slope for 35 m.

For a more detailed characterization of the behavior of hunting tigers and their prey, we will make use of specific examples.

On Januaryš 17, 1971, a hunt by the male "Emperor" tiger on wild boar, which had occurred along the right bank of Tigrovyi Stream in the basin of the Gornaya River, was written in their tracks.š The south-facing slope was of average steepness and was covered here by a Korean pine-oak forest that was almost without undergrowth and that therefore could be scanned from a great distance.š The depth of the snow cover was about 30 cm.š The hunt took place during the early part of the night, when, however, darkness was already complete, at a time when the wild boar were located in their "dens".š

In the valley of Tigrovyi Stream, the tiger turned away from the road, which had been laid down along Bol'shoi Stream.š Going upward along the stream, at first along its valley and then along its right bank slope, he approached a small, narrow side valley on his left.š After covering a distance of 390 m in his gradual ascent along the slope, he lay down for a time at the base of the trunk of an oak tree (lying on his stomach); 71 m beyond this, we found still another laying place (where the tiger lay on his side).š From there, the tiger abruptly turned to his left (relative to the previous direction of his movements) and began his ascent in a direction that was almost perpendicular to the dip of the slope.š Probably, the laying place at the oak tree reflected the first reaction of the tiger to the prey.š He had apparently scented wild boar that were located at a distance of 270 m from him (as measured in a straight line).š His scenting of the wild boar might have been facilitated by the air current, which is, as a rule, flowing downward toward the valley during the evening.š After 234 m of his ascent, the tiger once again lay down, being located now at a distance of 182 m from the wild boar.š Then, deviating somewhat to the right, he walked 132 m and again stopped.š A distance of 50 m still separated the tiger from the first two "dens" in which the wild boar lay.š This flanking maneuver, which had been carried out by the tiger's ascent along the slope, allowed the tiger to reach almost the exact same elevation on the slope as that of the wild boar.š Standing for a short time and then walking very slowly (each impression of the tiger's paws was covered with ice) for a further 32 m, he switched to leaping after his prey.

The tiger's charge followed when the tiger was at a distance of 18 m from the prey and also when the tiger was located somewhat higher along the slope.š One of the wild boar threw itself downward, but the other wild boar, the one that the tiger was striving to overtake, rushed along the slope, going upward.š At a distance of 58 m from the den where the tiger had frightened these wild boar, two more "dens" were located, in one of which a female wild boar was located, while three yearling wild boar rested in the other den, which was located slightly below it along the slope.š The tiger, running past the "den" of the solitary female at a distance of 5 m above her along the slope, rushed after her. Failing that, he then rushed after a piglet, which had run past him downward along the slope.š But the tiger also let him go by, and, turning about, the tiger fell down, after which he still attempted to continue the pursuit.š However, the tiger soon ceased his pursuit and, having made several strides at a trotting pace, he lay down.š The tiger experienced a lack of success, having attempted in the present instance to pursue various individual wild boar one after the other, and having rushed after the prey in all directions in a confused way.š All in all, the pursuit took place over a total distance of 560 m.

Figure 30. Tracks from a successful charge by a tigress in pursuit of an adult female wild boar (on the right: the leaps made by the tigress).

At a distance of 4 m from the tiger's laying place, where his hunt had ended, there was located yet another laying place where the tiger rolled on his backšfrom one side to the other, breaking a bush under himself in the process.š Still another short-term laying place was located 10 m from the previous laying place and lay along the tracks of a piglet that had run away.š From these laying places, the tiger headed upward along the slope at right angle to his original route.š Going 50 m farther, the tiger, having encountered the tracks of the other wild boar that had run away earlier, once again lay down, and this time, he was at his laying place for a much longer time.š Farther on, having walked for additional 390 m, the tiger stopped near a tree, trampled down the vegetation with his feet, and continued his path upward along the slope.š Small drops of blood remained in the tiger's tracks near the tree.š Such a flow of blood, as has already been mentioned, was observed quite often, its source being, apparently, cracks (due to chapping of the skin) in the fleshy parts of the soles of the tigers' feet.š After placing a great working load on soles of the tiger's feet (during the chase, etc.), these small wounds bled more severely.

One of the successful hunts made by the "Empress" tigress on wild boar, which took place under somewhat different circumstances than the hunt by the male tiger that we have just described, was observed by using the tigress' tracks on February 26, 1972, along the left bank slopes of the upper reaches of Bol'shoi Stream in the basin of the Orekhovka River.š The circumstances were different than in the previous case: a steep, north-facing slope that was covered with a Korean pine forest with an admixture of dark coniferous species.š The depth of the snow cover was 40-45 cm. The objects of the hunt were wild boar at the moment when they were feeding, probably, in the daytime.

The tigress emerged onto a small left tributary of the left bank slope of Bol'shoi Stream and lay down for a long time on a terrace of the small stream that had been warmed by the sun.š This laying place did not appear to be one used in a hunt, since the excrement of the animal (i.e. of the tigress) was discovered next to it.š Apparently, it was from this laying place that the tigress, which had rested there, had also discovered the wild boar.š Such an assumption is supported by the fact that she moved off from this site in a completely different direction than her previous one (up until this point, she had walked for more than 10 km along a path shared with a male tiger).

Having arisen from her laying place, the tigress headed upward along the terrace of the small stream and, 84 m farther on, she switched to making leaps.š After 78 m of her pursuit, she crossed the tracks of three wild boar that had rushed past her; she ascended farther along the slope at a walking pace.š The tigress walked 162 m from the site of her unsuccessful hunt and, turning aside to her left, the tigress reversed her course near a dead tree that had a cavern-lair in it (where she had stayed a year earlier with her cubs).š Having gone about 100 m downward along the slope, the tigress observed a piglet that had suddenly run toward the mountain and thus had approached the tigress.š The tigress made one spring in the direction of the piglet, then she covered 8 m by crawling.š After the piglet had come running up to the tigress, a spring by the tigress followed.š However, the wild boar succeeded in turning around and rushed downward along the steep slope.š The tigress succeeded in catching up with it only after pursuing the piglet for additional 28 m.

Having consumed part of the prey right where the hunt had ended, the female walked away for 10 m and lay down for a short while.š Then she walked upward along a small, narrow side valley and, having made a looped path of approximate length 260 m, she returned to the piglet along a wild boar path.š Having picked up the remains of the prey, the tigress hid them under a cover of coniferous trees.š Judging by the skull that we examined, the piglet weighed not less than 30 kg.š The path made by dragging the prey was directed straight toward the mountain and extended for 97 m.š At a distance of 40 m from the piglet, which had now been left in a new location, a shelter was located underneath an overturned tree, this shelter having been repeatedly visited by this tigress in the past.š Apparently, this was the reason that she had dragged her prey over there in order to have it closer to her shelter.š She walked from the prey to the lair 3-4 times, having tramped down a track.š Once, returning to her prey, the tigress made a spring when she was halfway to the prey, clearly with the aim of scaring away crows.š She spent 2-3 days near the prey.š At a distance of 20 m from the shelter, we found excrement from three different defecations as well as two urine spots (a third urine spot was found close to the prey).š Only the head, the wrists, the feet, and the torn up skin of the piglet's carcass remained.š From the site of the hunt, the tigress once again headed along the tracks of the male.

The next example of the tigers' hunts on wild boar that has been selected for a detailed description was distinguished by the fact that the hunt was accompanied by extremely difficult conditions, since a great deal of snow was present.š The hunt took place on February 24, 1973, on the left bank slopes of Pechenyi Stream in the basin of the Malinovka River.š The site of the hunt was a west-facing slope of medium steepness, where the depth of the snow cover reached 60-65 cm.š The attack on the wild boar, which were located in their laying places, took place during the dark hours.

At Pechenyi Stream, the "Miniature" tigress turned off the road that had been laid down along the valley of the Malinovka River.š Her path upward along the stream ran at first along the channel of the stream, which was free of snow, and then her path ran along the riverine forest, where she kept to trails made by wild boar.š Here the tigress encountered a piglet that had perished from starvation after it had lain down in the "den"; the tigress had dragged the piglet 8 m to one side, but she did not begin eating it.š She abruptly turned from the site of her discovery in the direction of the channel of the stream and, emerging onto its left bank slope, followed the fresh tracks of wild boar. After the tigress had followed these tracks for a distance of 490 m, they led her to three wild boar, which were lying along the slope.š The hunt began "from the tigress' path".š When there remained a distance of only about 20 m to the "den" that was nearest the predator, a wild boar (a piglet) jumped out of the den and rushed downward along the slope in a direction perpendicular to that of the tiger's approach.š He ran away along numerous frozen ditches made by wild boar, in an arc to the right, but the tigress, pursuing him, "cut off" this arc.š The piglet could not run away along a straight path, since deep snow stood around the continuous ditches like a wall.š After going 338 m, the piglet jumped back onto the trail, along which the tigress had approached, attempting by making a circular path to reach the "dens", around which the snow had been well trodden down and dug up.š But the predator sent him downward along the path and, after 52 m of pursuit, overtook the piglet.š All in all, the pursuit continued for a distance of about 390 m.

Having fed for a while at the kill site, the tigress dragged away the remains of the piglet along the track that she had made during her approach.š After having brought the prey over a distance of 78 m to an old "den" of the wild boar, the tigress once again ate for a while, after which she left to rest at a site, which was located approximately 90 m higher up on the slope, where the wild boar had been frightened off.š The path between the laying places and the prey was well trodden.š Later, the tigress brought the remains of the piglet to her resting site.š The tigress spent not less than 24 hours near her prey.š The remains that she abandoned were in approximately the same state as those in the previous case.š Six urine spots were noted along the track between the prey and the tigress' resting site.š Excrement was found along a 290 m-long stretch of the tracks left by the departing tigress.š

Hunts on Manchurian Deer:

Tigers most often hunt these ungulates in the forests of river valleys.š Of the 38 hunts on Manchurian deer that we recorded, 22 hunts (57.9%) took place in floodplain habitats and on terraces located above these floodplains.š In eight cases, the tigers attacked their prey from roads that had been laid down along the valleys.š Manchurian deer often forage along the sides of roads, and the predators can approach them along the tramped down roadbed without making a sound.š For comparison, we note that a hunt on wild boar in which the tiger began his hunt from the road was observed only once--in a year that had poor crops of the main foods of the wild pigs.

Hunts on Manchurian deer took place, as a rule, during the dark portion of the 24-hour period [in 16 cases out of 20 (cf. Table 15)].š In six instances, these were pre-dawn hunts, and, in five cases, the hunts occurred during the evening hours.š If the phases of activity and rest during the course of the 24-hr cycle are in general opposite for the tiger and the wild boar, then the 24-hour activity rhythm of the Manchurian deer is similar to that of the tiger.š Manchurian deer spend the greater part of the daylight hours in their laying places, and they feed most actively with the arrival of dusk and during the pre-dawn hours.š Accordingly, Manchurian deer become the objects of the tigers' hunts while foraging more often than do wild boar (cf., Table 15).

The minimum, maximum, and mean distances of discovery of Manchurian deer by a tiger (for 18 hunts) equaled 40 m, 320 m, and 138 m, respectively.š The predators found these ungulates, as a rule, at a somewhat shorter distance than the distances at which they discovered wild boar.š The latter species creates much more noise when on its feeding grounds, especially when the wild boar are present in a herd, than do grazing Manchurian deer.š Tigers can determine the locations of wild boar by the sounds that they emit during the period of the rut at a distance of a kilometer or more (during the roaring period, Manchurian deer obviously attract the attention of predators to themselves at a still greater distance, however, we did not conduct observations during autumn).š It is easier to discover wild boar not only by sound, but also by their odor, which is clearly stronger (than that of Manchurian deer) and which diffuses farther.

The association between methods of hunting used by tigers also varies in relation to their main prey species.š Manchurian deer are more rarely attacked "from the tiger's path" (11 of 34 cases--32.4%).š The great cautiousness of Manchurian deer in comparison with wild boar makes them less accessible prey for the predators, in connection with which, when hunting on deer (Cervidae), tigers most often have recourse to a long period of stalking with various kinds of interruptions (stops, laying places used during the hunt).š Hunts that included such elements were observed 19 times (55.9%).š Tigers utilized evasive maneuvers almost three times less frequently than during their stalking of wild boar, but attempts to approach the prey from short distances by crawling were more frequent.š A hunt from an ambush was observed in 4 cases.

Tigers began making leaps after Manchurian deer from a distance of 3 m up to 80 m (cf., Table 16).š The mean distance of approach to Manchurian deer when the tigers were stalking their prey was 25.6 m (for 24 hunts).š The length of the pursuit ranged from 10 m up to 260 m, on average, 63 m (for 26 hunts).š Without the inclusion of two deviant cases involving the pursuit of Manchurian deer at distances of 200 m or more, the mean value is 47 m.š Thus, the tigers' pursuit distance when after Manchurian deer is usually shorter by a factor of two as compared to their pursuit of wild boar.

The positions of predators and prey relative to the topography are in approximately the same relationship as with hunts on wild boar.š In 11 cases (47.8%), tigers were situated above the prey; in three cases where tigers attacked, the predators were below the prey (13.1%), and tigers were at the same elevation as the prey on 9 occasions (39.1%).

In the majority of cases, the tigers killed the Manchurian deer that they had overtaken right on the spot.š A struggle involving the predator being "dragged along" by the prey was observed in two instances when the prey was adult deer, and once during an attack on a yearling male deer (or "shpil'nik").š The tiger finally succeeded in stopping the adult animals after the tiger had been dragged for a distance of 5 meters in the first instance and 55 meters in the second case.š The yearling male deer was stopped after dragging the tiger for 10 m.

We now present detailed descriptions of several specific instances of hunts by tigers on Manchurian deer.

On March 13, 1971, along the upper course of Levyi Igristyi Stream (in the basin of the Gornaya River), the "Miniature" tigress attempted a daytime stalk of Manchurian deer that she had discovered at the deers' laying sites.š The site of the action was a Korean pine-deciduous forest (which had been thinned by logging) on the terrace of the stream.š The depth of the snow cover was about 50 cm.

The tigress, having ascended the valley along the foothills of the left bank slope, suddenly abruptly turned to her left, crossed the floodplain of the stream and its channel, and emerged onto a right bank terrace.š Continuing her path across the valley, she walked for approximately 360 m and lay down on a small snow-covered knoll, lying slightly toward her right side.š Within a distance of 260 m from this site, we found two of her short-term laying places that followed one after the other.š There were many tracks of Manchurian deer here as well as the laying places of these deer.š The tigress walked for another 200 m, abruptly turned to the right, and, after taking several steps, she lay down, resting on her thorax and folding her rear legs underneath her.š From this site, she apparently scented the Manchurian deer that were located on their laying places at a distance of about 200 m from the tigress.

The tigress moved toward them, having left two similar laying places right next to each other on the snow cover over a distance of 32 m.š Farther on, have decreased the distance between herself and the deer to 65 m, she stopped and kneaded with her feet in one place for a long time, after which her charge followed over a distance of 10 m.š Having made 20 leaps to the laying place, from which she frightened off five Manchurian deer, the tigress ceased her pursuit and went along at a walking pace.š The sprint was made at a distance of 55 m from the deer.š The frightened deer traversed a distance of 250 m by leaping, with some leaps reaching 7 m in length, and after that they switched to a walk.š The leaps made by the predator were, from the very first, short, as if the tiger were hesitant; we felt that the animal did not exert all its strength during the pursuit.š Without even knowing the result of the hunt, it was possible to assume from these tracks that the hunt would finish as an unsuccessful one.š After her attempt at pursuing the Manchurian deer, the tigress returned to her original route from which she had deviated after having begun the stalk, and she move farther on in her previous direction.

The male "Emperor" tiger hunted a yearling Manchurian deer (a shpil'nik) on January 27-28, 1972, at the sources of Bol'shoi Stream (in the basin of the Orekhovka River). The entire hunt took place along a north slope of medium steepness, which was covered with a Korean pine forest that had a significant admixture of dark coniferous species.š The depth of the snow cover here reached 45 cm.š The hunt probably took place during the dark portion of the 24-hour period.š The Manchurian deer was feeding prior to the tiger's attack on it.

The tiger, having ascended the slope obliquely up to the divide that stretched between Malyi and Bol'shoi Streams, abruptly turned toward the side nearest the crest when he was at a distance of 13 m from the mountain ridge.š From here, the tiger heard a Manchurian deer that was located on the opposite side of a small spur, at a distance of 110 m below the line of the divide of the watershed.š Having crossed over onto the slope where the Manchurian deer was grazing, the tiger slowly went downward, deviating from a direct descent by going somewhat to his left, not making any prolonged stops and not lying down during his descent.š At the same time, the Manchurian deer gradually moved upward along the slope.š The tiger's charge followed at the moment when the distance between them had decreased to 11 m.š The Manchurian deer at first made one leap to the side in order to make an 180o turn, and, having turned about completely, rushed downward along the slope.š But 19 m farther along in the tiger's pursuit, the tiger already "was hanging" from the body of the Manchurian deer.š While restraining the prey, which was continuing to run, the tiger left impressions of his tail to one side in the snow.š Having made four leaps from under the predator, the Manchurian deer fell down and was no longer able to get up from that position.

The tiger dragged the prey for a distance of 43 m downward along the slope and placed it at the base of a sprawling Korean pine, which stood next to a heap of trees, where he ate it for a while.š Having assuaged his hunger, the tiger went to seek a shelter.š He investigated an overturned tree, which was located downward along the slope, but he did not lie down here.š Then he began to ascend in the direction of the divide of the watershed, and, leaving his prey while walking a distance of 78 m, he stopped, having placed his front paws on a fallen tree trunk.š Two roe deer that were 100 m downward along the slope attracted his attention.š Stealing up to them, he switched to making leaps.š His charge at the prey occurred when a distance of 16 m remained between the tiger and the roe deer, but the tiger was forced to run upward along the slope.š The pursuit turned out to be unsuccessful.š Terminating his pursuit at the crest of the divide after going 24 m, the animal returned to the prey that he had killed earlier.

The tiger lived for approximately three days near his prey.š The lair on which the tiger most often rested was located at a distance of 25 m from the location of the tiger's feeding site and was located upward along the slope on the ground near the root of an overturned tree.š Another long-term unsheltered laying place, which the tiger, apparently, used during the daytime, was situated at a distance of 13 m from the remains of the prey along the tiger's path to his lair.š There was also a similar laying place lower down along the slope at a distance of 50 m from the prey.š In addition to those that have already been mentioned, still another two laying places (where the tiger lay on his side) were found next to the Manchurian deer as well as one short-term laying place found under a group of fallen trees.š The predator's excrement was left in four places: two sites were near the lair and two were located along his tracks at a distance of 50 m from the prey.š

In all instances, the excrement was fluid, black in color, and of a shape reminiscent of a flat cake.š There were five urine stains on the snow here, among them four located near the lair.š All that remained from the prey were: the head with the neck, the rear feet below the knee joints, and the partially consumed front legs.

The following example involves a hunt conducted by a tiger where the prey was discovered from the roadbed.š On February 24, 1971, the male "Emperor" tiger killed a Manchurian deer in a section of floodplain forest in the valley of the Gornaya River that was situated at a distance of 45 m from an actively used logging road.š The depth of the snow cover in the forest was 30-35 cm.š The hunt took place in the pre-dawn hours.

The tiger, having walked along the road, suddenly made a 90o turn to the right of his path, apparently, having heard a Manchurian deer that was located along his path at a distance of approximately 60 m from him. The Manchurian deer (this was an adult female) was feeding on water plants while standing in an unfrozen channel that was about 4 m. in width.š The bank of the stream that was closest to the tiger was gently sloping, but the other shore was precipitous.š Bearing somewhat to his right and moving away from the road, the tiger crossed the channel downstream from the Manchurian deer, and, having made an arc to his left, he crept up on his prey, which was hidden from him by the shoulder of the steep bank.š The entire path of the predator in this flanking maneuver totaled 470 m.š The back of the Manchurian deer, which was standing in the channel at a distance of 2-3 m from the ledge of the stream bank, probably protruded somewhat above the ledge.š A leap by the tiger from the shore toward the back of the prey followed.š The Manchurian deer, which was killed on the spot with a strike by the tiger's canine teeth in the neck region, remained lying in the channel.š The upper part of the carcass projected from the water, where the tiger was also located while eating the carcass.š However, the tiger ate only a small amount--only part of the "flesh" of a rear leg--after which he went 30 m away toward the road and lay down there.š When he arose from his laying place, he emerged onto the road, went several steps along it to the right, and turned back toward his prey.š But he no longer touched his prey; instead, he headed across the valley of the stream.š The prey was abandoned, probably because of the nearness of the logging road.

* * *

Tigers generally hunt other species of ungulates, in particular roe deer, the same way as they hunt Manchurian deer.š These predators do not make a special attempt to pursue musk deer: they become the prey of tigers only as a result of a chance encounter.š For example, in one instance a musk deer was killed after it had unexpectedly sprung out by a tiger that had already passed by.

We encountered bears (a female Himalayan bear with a cub) among the tigers' prey only once: they were killed by a male tiger in an above-ground den, which was situated under an overturned tree at the root end of the tree (Fig. 31).

Tigers' encounters with dogs in the taiga occur mainly during the commercial hunting season.š It is the dogs themselves, as a rule, that, having gotten carried away by excitement of the hunt, approach the tigers to within a dangerous distance and become their prey (Fig. 32).š It was in exactly this manner that a hunt by a tiger on a dog once took place as we later determined from our tracking.š In two other cases, tigers attacked dogs which had wandered far from settlements without their owners and which had attempted to pursue ungulates.

One such hunt, where a charge by the tiger ensued when the tiger was located at a distance of 53 m from the dog, yet where the tiger's pursuit of the dog stretched for 154 m, ended with a lack of success.š In another instance, a tiger searched out a dog during the night by using its tracks, which stretched from a road into the forest.š Having covered a distance of 1.8 km with his "tracking", the tiger approached to within a distance of 30 m of the dog.š The tiger approached at his usual pace, without using laying places and without apparently stalking the dog.š The dog had lain down under a spruce tree (Picea spp.).š Having been startled, the dog rushed toward the road; but the tiger, having cut a corner, reached the dog after covering 66 m (as a series of 19 leaps).š The leaps made by the dog did not exceed 2 m in length.

The tigers' methods of killing different species of prey can only be evaluated in rare cases.š We succeeded in discovering wounds on the necks of animals that had been killed.š In the case of yearling wild boar, the cervical vertebrae were sometimes crushed.

Figure 31. An above-ground den of a female Himalayan bear with her cub.

Figure 32. The remains of a dog killed by the "Miniature" tigress.š

Success Rate of Hunts and Factors Influencing Hunting Success:

It is already apparent from the examples presented above that the success of hunts made by an attacking tiger depends above all on the distance to which the tiger succeeds in approaching the prey, in other words, on the suddenness of the charge.š However, other factors also play an essential role: the location of the predator above or below the prey, the depth of the snow cover, various types of unexpected events either favoring the hunt, or, on the contrary, complicating it.š With regard to the positions of the animals with respect to the topography, tigers more frequently overcome prey if they are situated above the prey or at the same elevation before the attack than when charging the prey from below.

The mean distance at which the charge of the tiger began is shorter by a factor of two (or slightly greater than two) for successful hunts versus unsuccessful hunts.š If we examine only hunts on Manchurian deer, the difference increases to almost a factor of three (cf. Table 16).š Real chances for successful hunts appear only when the tiger succeeds in approaching a wild boar to within a distance not greater than 30 m, and a Manchurian deer to within 20 m (Yudakov & Nikolaev 1977).š But charges from such distances also frequently turn out to be unsuccessful.š In one case, the attack turned out to be unsuccessful even with an approach to within 4 m.š The tiger was hindered by the presence of a Korean pine tree that concealed a wild boar that had already begun to escape from the tiger's pursuit; the predator's charge, moreover, was directed upward from a position below the prey.š In sum, approximately 75 % of all successful hunts (both on wild boar and on Manchurian deer) take place in those situations where the tigers attacked their prey from a distance of not less than 15 m from the prey.

Differences in the mean distances for the start of the tiger's charge in comparisons of successful hunts on animals of various ages are clearly revealed only in the case of wild boar (cf. Table 16).š When capturing young animals, it is sufficient for the tiger to approach the prey to within a distance 1.5 times greater than that required during the stalking of adult animals of this species.

Cases where the tiger succeeded in seizing the prey on the spot, without first pursuing it, occur as rare exceptions.š Even Manchurian deer and, in one instance, a roe deer were killed with a single spring; one Manchurian deer and one roe deer were seized after several leaps by the predator without the prey having succeeded in even moving from the site.š As a rule, the tigers' hunts cannot avoid including a short chase in pursuit of the prey (cf. Table 16).š The length of the pursuit in approximately 70% of all successful hunts does not exceed 60 m for wild boar or 40 m for Manchurian deer.š Having overtaken wild boar, tigers sometimes run after them for 150 m, 390 m, and even 900 m.š The outcome of hunts on Manchurian deer is determined at shorter distances: a pursuit over a distance greater than 75 m was not successful on even a single occasion.

In addition, the minimum pursuit distance that brings tigers success is greater for Manchurian deer than for wild boar: tigers more often catch up with the latter species at shorter distances.š The pursuit of adult wild boar lasts longer than does the pursuit of yearling wild boar, which is reflected in both the mean and the minimum values (cf. Table 16).š Analogous differences appear in an even more marked form with respect to adult and young Manchurian deer where the rates vary by a factor of two.š As can be seen from these data, the pursuit of young animals involves the tiger in a significantly lower energetic expenditure in comparison with a hunt on adult individuals.

In the case of unsuccessful hunts on wild boar, the length of the tiger's path during a pursuit reaches half a kilometer; on average, it is 200 m.š Manchurian deer are usually not pursued by tigers over distances greater than 100 m.š A strenuous charge by the tiger sometimes changes to an unhurried walk during the pursuit of the prey along the prey's tracks.š In one case, the tiger followed a herd of wild boar along their trail for a distance of 1700 m, and, in another instance, when there were three wild boar, the distance was 2500 m, and here this "tracking" ended in an attack by the tiger.š Similar pursuits of Manchurian deer continued in two cases for 1200 m and 2200 m.

An approach to the prey does not always end in an attack.š If the prey discovers the predator sooner than a charge that can be relied upon to succeed becomes possible, the hunt is usually interrupted at the stalking stage.š This was true of 5 out of 18 cases of unsuccessful hunts on wild boar (27.8%) and of 9 out of 24 unsuccessful hunts on Manchurian deer (37.5%).š For roe deer, the percentage of such cases increases to 50%.š Cases involving the stalking of Manchurian deer without the tiger completing a final charge are observed more frequently when deep snow is present.š Sometimes, the hunt is also interrupted from another cause--from the readiness of the prey to exhibit resistance.š In one case, a male tiger decided not to attack a large wild boar.š Having discovered the latter animal in its "den", the tiger made eight leaps in the direction of this animal, but the tiger stopped when he was at a distance of 30 m from the boar, and then moved at a walking pace.š This animal was, in all probability, a young male boar, which, having arisen from the "den", took up a defensive posture, which forced the tiger to withhold his attack.š The predator was forced to turn aside, after which he left that place.š The case just described is quite reminiscent of an unsuccessful hunt of a tiger on a Gaur bull, which was observed in India by G. Schaller (Schaller, 1967).

Once during tracking we also noted the following fact: a tiger retreated, not attempting to hunt a large male brown bear in its den.š The bear had set himself up for the winter in a small depression that he had dug near a fallen, broken-off shrub (Fig. 33), and the bear was quite visible. The tiger, having encountered the bear by chance, abruptly turned around at a distance of more than 25 m from this place and walked in the opposite direction by following his own old tracks.

No essential differences appeared in the outcomes of hunts on foraging animals in comparison with hunts on animals resting in their laying places.š The success of a hunt depends to a much greater degree on whether the tiger is stalking a single animal or a group of animals (Table 17).š A hunt on Manchurian deer that had gathered in small groups was, as a rule, unsuccessful. The tigers' prey consists mostly of solitary animals that discover the danger at a later time.š For wild boar, the differences are in the opposite direction and are not so marked.š It is mainly adult animals, particularly male boar, that live as solitary animals: they are more cautious and a hunt on them is accompanied with greater difficulties than is the pursuit of young animals.š Wild boar in a herd not only make a lot of noise during feeding, but also disperse widely, and when alarmed, sometimes scatter in different directions, which facilitates the approach of the tiger to a potential prey.š Piglets and gilts in search of the herd at times return to the very same site from which they had been frightened away by the tiger.š Once, a predator, which had been resting after an unsuccessful hunt in a "den" that had been deserted by the wild boar, was closely approached by a yearling wild boar, which also became the tiger's prey.

The success of the hunt also depends on the condition of the snow cover, above all on its depth.š Both for the prey animals and for the predator, travel across snow is accompanied by an additional expenditure of energy, and by a decreased running speed.š As G. F. Bromlei (1959) has shown, a judgment on the degree to which snow serves as an obstacle for the movement of different animal species can be made using the ratio of the weight loading on the animal's tracks to the chest height of the animal.š The dimensionality of this index is: g/cm2/cm.š The loading on the footprints of a Manchurian deer is, on average, 675 g/cm2; for wild boar, it is 414 g/cm2 (Bromlei 1970).š However, the index measuring the difficulty of movement in snow for the Manchurian deer is 8.6 g/cm2/cm, considerably less than that for wild boar: 10.0 g/cm2/cm.š Variation in snow depth creates conditions that also act unequally on animals of different ages that belong to a single species.š Thus, in the case of wild boar, the weight loading on the animal's footprints changes with age from 184 g/cm2 to 450 g/cm2, with seasonal fluctuations also having major effects (Bromlei 1964).š The loading on the tiger's footprints is, according to various sources, in the range of 149-158 g/cm2 (Formozov 1946, Bromlei 1970).š According to our data, it is (n=8) on average, 226 g/cm2, and the index of difficulty of movement on snow is 4.3 g/cm2/cm.š This is less by a factor of two than even the value for Manchurian deer.š

Figure 33. The den of a large male brown bear ("sidun").

Table 17.

Success of tigers' hunts on solitary animals and on groups of animals.


Solitary Animalsššššššššššššššššššššššššššššššš Groups of Animals
Object of the Tiger's Hunt Total Successful Unsuccessful Total Successful Unsuccessful


No. Hunts Abs. % Abs. % No. Hunts Abs. % Abs. %
Wild Boar 11 3 27.3 8 72.7 26 15 57.7 11 42.3
Manchurian Deer 23 8 34.8 15 65.2 13 1 7.7 12 92.3

When the snow cover is greater in depth than 50 cm, the success rate of hunts made by tigers on wild boar in conditions of deep snow increases by almost 1.5 times in comparison with hunts made when little snow is present.š In 1972/73, which was a season of much snow, yearling wild boar turned out to be almost without any protection from tigers.š The predators overtook wild boar on their trails and sometimes caught a piglet that was lagging behind the herd without even having to make a spring toward the prey.š Hunts on Manchurian deer have a success rate that changes hardly at all in relation to the depth of the snow cover (according to our data, in deep snow, it even decreases by a factor of 1.1).š Manchurian deer possess such a clear overall advantage in running speed that tigers do not always decide to begin pursuit of this deer even from a distance of only 10-15 m.š Once, when the depth of the snow cover was 60-70 cm, a tiger, which was in an ambush position, "allowed" a Manchurian deer to pass by him at a distance of only 13 m.

The leaps of an attacking tiger are approximately the same length as those of wild boar, but the leaps of Manchurian deer are considerably longer (Yudakov & Nikolaev 1977).š The initial spring of a tiger toward the prey did not exceed 3 m in even a single case; often, this spring turns out to be the shortest one in the series of leaps involved in a single pursuit.š This spring was not longer even in those cases when, in a successful attack, this leap turned out to be the only one.š The mean length of the initial leap was 1.8 m.š Rushing after prey, the tiger leaps on average a distance of 3.4 m, whereas a wild boar that is escaping from the tiger leaps on average 3.1 m., and an escaping Manchurian deer leaps 3.9 m on average.š The maximum length of the tiger's leaps (among those that we measured) equaled 6 m; it was directed down a slope. The longest leaps made by wild boar and by Manchurian deer reached 5.5 m and 7 m., respectively.

1The data were collected by us in collaboration with V. G. Yudin with the support of the Regional Directorate of the Hunting Industry of the Primorskyi Regional Executive Committee.

Manchurian deer estimate very precisely the distance that they need to run (while escaping): they switch to an unhurried walking pace when they are 200-250 m from the site where the tiger stopped pursuing them.š The following case is very illuminating in this regard.š The tiger, having approached to within 28 m. of two Manchurian deer in their laying place, attempted to attack them, but after 48 m of pursuit, the tiger turned around and walked backward along his previous route; the deer themselves, having run only 165 m from this site, stopped and again lay down.š Wild boar behave differently in similar situations: they rush away leaping sometimes for more than a kilometer from the site of tiger's attack, and even after this, they remain on guard.

Measurements of over 500 leaps (for all hunts) established that the mean length of a tiger's leap during successful hunts is somewhat greater (by 0.2 m) than during unsuccessful hunts.š The difference in the mean length of leaps for a shallow depth of snow cover and for deep snow is, as has already been noted (Matyushkin & Yudakov 1974), approximately 0.5 m.

Summary values measuring the successfulness of the tigers' hunts on wild boar and on Manchurian deer are different for the two species.š These predators capture wild boar in more than half of their attacks (54.5% for 33 hunts), but they overtake Manchurian deer in less than a third of their charges on them (28.9% for 38 hunts).š For the total aggregate of all the data that we have collected, including data from other regions in Primorskyi district, these indices are 51.3% (n=39) and 30% (n=40), respectively.

Utilization of the Prey:

The tiger eats the prey at the site where it was killed or drags it, and later, its remains, to a new site.š Sometimes, the tiger drags the carcass in several stages, partially consuming it at the various stops.š The prey is transported by pulling it along, in very rare cases with the prey hanging from the tiger's jaws.š In 18 (47.4%) out of 38 cases, the prey was fed on for a while at the kill site, but in 20 cases (52.6%), the prey was dragged for distances ranging from 13 m to 650 m.š In 9 cases (45%), the trench extended for a distance of 13 m to 30 m, in 5 cases (25%) for distances ranging from 30 m to 50 m and in 6 cases (30%) for distances greater than 70 m.š Most often, tigers drag the prey up to a "den" made by wild boar, which serves them as a resting site, or to some sort of nearby shelter, for example, under the hanging branches of coniferous trees (Fig. 34).š These searches for a more secluded site have the aim, apparently, of masking themselves and the prey from importunate crows.š Sometimes, the tigers move their prey to comfortable lairs if such lairs are present nearby.š When the need arises for the tigers to remove themselves from a source of disturbance, for example, from parts of a road that are suitable for vehicles, the length of the trench can be greater.š Animals that have been killed on slopes are usually transported downward along the slope by the tigers.š Adult wild boar and adult Manchurian deer are most often eaten on the spot (i.e., at the kill site).

The series of footprints of a tiger that is dragging prey almost always extends to one side of the trench.š In only one case did the series of footprints lie along the trench, when the tiger dragged a yearling wild boar along in front of himself, apparently holding it by the neck and almost lifting up the anterior portion of the carcass.š Tigers prefer to drag large animals (cows, horses) backwards by holding onto them by the neck or by the rear portion of the carcass.š Indications encountered in the literature that Amur tigers can carry large animals and can even leap while holding them, we consider to be without foundation.š According to our observations, transport of the prey while it is hanging from the tigers' jaws is generally uncharacteristic of tigers.š In connection with this, it is necessary to present information on four cases where such behavior nevertheless took place, having noted beforehand that whole carcasses of wild boar and Manchurian deer were not once carried in this manner.

Once, having killed a roe deer, a very large male tiger (with a "heel" width of 13.5 cm) carried it in its jaws.š This was the heaviest load carried by a tiger among those that we observed.š Furrows that had been left in the snow by the hanging legs of the prey stretched along both sides of the predator's tracks (Fig. 35).š In total, the tiger carried the carcass about 300 m; along this path, he stopped twice, placing the carcass on the snow.š The prey was transported from a south-facing slope to a northerly slope.š In another case, a tigress, having killed a musk deer, walked with it for 55 m, sometimes lying down and sometimes stopping along the way.š She dragged it along, making trench in the snow, or carried it in her jaws; in the latter case the legs of the prey also created furrows in the snow.š At the laying place located at the end of this section of the tiger's path there remained a lot of musk deer hide that had been ripped off by the tigers' claws.š Later, the prey was transported 20 m farther, and it was consumed at the base of a Korean pine tree (Fig. 36).š In the next case, a dog that had been killed by a tigress later had its remains dragged for a distance of 650 m.š The predator with her prey went to hide behind a small mountain spur to avoid a human that was present in the vicinity.š Having dragged the prey for the first 260 m by making a trench, the tigress stopped and ate some of it, dragged it for a further 195 m, and then, for a distance of almost 200 m, the tigress carried the remains of the dog in her jaws.š Finally, a tigress once dragged the remains of a yearling wild boar over a distance of 170 m, a case that we have already mentioned.š She first fed for a while on the piglet that she had killed at the kill site, then, while dragging the prey over a distance of 78 m, the tigress continued her meal; next she rested in a wild boar den, and from there, the tigress carried the remains of the piglet in her jaws for a further 92 m.

Figure 34. Prey--a Himalayan bear--concealed by the male "Emperor" tiger under the hanging branches of a spruce tree (Picea spp.).

Figure 35. Arc-like furrows in the snow--tracks from the hanging legs of a roe deer that was transported in the jaws of a male tiger.

These predators usually begin to eat the prey beginning with the rear part of the thorax and the legs. In three instances, the tigers first removed the skin from the carcass with their claws (Fig. 37).š The prey is eaten together with the skin or else individual pieces of skin that have been bitten off remain.š In the case of large prey, the tubular bones can also be consumed, or they can also be left behind as gnawed bones.š The internal organs of recently killed prey are sometimes pushed to one side of the carcass by the tigers (Fig. 38). Spots of blood on sites where the prey has been eaten were found only very rarely, as is also true for blood spots on the tracks of tigers moving around prey.

The selection by tigers of rest sites near their prey is determined by the presence of convenient lairs, wild boar "dens", and areas that are accessible to solar rays at such locations.š From one to four long-term lairs situated in different places can be located in the vicinity of the prey.š Their number and location depends on the length of time that the predator has spent there and on the conditions present in each specific area.š Laying places of tigers near killed wild boar were observed both alongside the prey as well as at a distance of 92 m from it, on average at a distance of 22 m from the prey.š The most distant laying places are located in lairs and in the "dens" of wild boar, if such places do not turn out to be located in the vicinity of the prey.š In half of all cases when wild boar was the prey, the predators utilized the "den" of the wild boar as a long-term lair.š Tigers lie down to rest at significantly greater distances from Manchurian deer on which the tigers have fed.š In part, this is because comfortable sites for lairs were not available near this type of prey.š The most distant of the lairs from a site of successful hunt on Manchurian deer was located 250 m from the prey, once again in the "den" of wild boar.

During the daytime, apparently, the tigers more frequently rest in the immediate vicinity of the prey, but at night, the tigers rest in a shelter, if such shelters exist nearby.š When the tiger has spent a long time near the prey, a well-trampled path extends from the resting lair to the meal site.š Sometimes, tigers unexpectedly change over to leaping while on their way to the remains of their prey; apparently, the tigers are scaring off crows.

"Tigers spend from several hours to 3-4 days near their prey...The utilization of the prey by the tiger depends not so much on the dimensions of the prey, so much as on the degree of satiation of the tiger.š Quite often, adult males will consume not more than 25-50% of an animal that they have killed" (Yudakov 1973, p. 93).š In eight (20%) cases, not including three instances when the tigers were frightened away from their prey during our tracking, the killed animals were only half-consumed (or even less) (Fig. 39).š Three such corpses were found near a road used for transport.š One corpse of a Manchurian deer had approximately 5% of it eaten, a second had 20% eaten, and a third was half-eaten.š The fact that the animals had been disturbed, of course, influenced the degree of utilization of the prey.š This is particularly relevant to the first case--the prey had been killed in the early morning prior to the start of motor transport.š The two other hunts occurred in the late evening, and the tigers moved away from their prey at an unhurried walking pace.

The remaining five cases, when the predators had not been alarmed, but the prey, nevertheless, remained to a significant extent uneaten, can be dealt with from the position of a hypothesis of "surplus killing".

Hunts by tigers that followed immediately upon the completion of a successful hunt can serve as support for this idea.š In one such hunt, a tigress had killed a roe deer with the tigress having spent not more than 3 hours near the dead prey and having almost completely consumed the prey.š After going 1.2 km from the site of the successful hunt, she already attempted to pursue a wild boar, but unsuccessfully. Yet, once again, 1.3 km farther on, she killed a yearling wild boar and consumed almost half of it.š Between these two prey kills, she had only a single laying place, which she used briefly.š In another instance, a male tiger hunted roe deer in the vicinity of a Manchurian deer that he had partially consumed.š Finally, once a tigress, having killed one of the three roe deer that she had pursued, continued her pursuit of the two remaining roe deer.š Here, the roe deer was killed as though in passing, without a stop or any interruption in the tigress' galloping motion.š The pursuit continued for 95 m and ended unsuccessfully, after which the tigress returned to her kill.š

Figure 36. The remains of a musk deer killed by the "Miniature" tigress.

Figure 37. A large male boar (in the background) that had been killed by a male tiger.š Pieces of boar skin that have been removed from the boar by the tiger lie in the foreground.

Figure 38. The remains of a yearling Manchurian deer that was consumed by a tigress with her two cubs.

Figure 39. A yearling wild boar that had been killed by the "Empress" tigress and which remained uneaten.š

Tigers also behave in a similar way during an attack on calves of domestic cattle.š We happened to observe the results of such a hunt on September 11, 1969, in the vicinity of the settlement of Sokol'chi (in Lazovskii district), when a tiger immediately killed three animals from among a herd consisting of young animals, which all had rushed from the enclosure when the predator attacked. The dead prey lay at distances of approximately 100 m from each other in a pattern that almost resembled the three points of a triangle.

The return of tigers to the remains of prey that they had killed earlier was observed in only 4 instances.š Once, a female with cubs came up to the remains of a bear and dug the remains out of the snow, where they had been concealed by an observer; later she visited these remains by herself (i.e., without her cubs).š This same tigress returned to the remains of a Manchurian deer that she had killed.š Only one similar case was recorded for male tigers.

Females with cubs and solitary tigresses consume their prey, as a rule, almost completely.š Only the skull, the legs (below the tarsal and wrist joints), part of the intestines and pieces of the skin are left.

Feeding by tigers on carrion, which was very rarely observed in the study region, took place during a period when a great deal of snow was present.š In two cases, females fed on dead roe deer, and once a tigress ate a bait that was located near a steel trap.š Twice we observed that tigers did not start to consume Manchurian deer that had died of starvation that the tigers had discovered.

Copyright A. G. Yudakov,I. G. Nikolaev

Copyright K. Lofdahl, A. Shevlakov, 2004 (English translation)