Chapter 11:
Marking Behavior: Forms and Frequency of Appearance in Different Contexts
Traces of urination and defecation are very frequently encountered along
tiger tracks, sometimes clearly more often than might be expected in relation
to the simple physiological necessity of the evacuation of the tiger's urinary
bladder and large intestine. As is well known, urine and fecal matter acquire
significance for neighboring individuals as chemical signals, which carry
information, particularly concerning territorial occupancy, the readiness
of the female for mating, etc. The marking function of urine and feces
is clearly borne out by the following observations: 1) tigers commonly apportion
urine or feces in small amounts over short intervals of distance (or time);
2) the action of chemical signals is amplified by visual signals--sites
that are marked receive attention, contrasting with the background surrounding
them; 3) when approaching marked objects, the animals change the direction
in which they are walking, sometimes rather abruptly.
Most often, tigers leave urine marks on objects having a flat, vertical
surface: rocks, tree trunks, or isolated bushes. Urine spots on the horizontal
surface of the snow are found over 10 times less commonly (vs. marks on
vertical surfaces) in males but approximately 4 times less often in females
(Tables 10 & 11). It is
possible to form a judgment about the behavior of tigers during their marking
of flat, vertical surfaces from observations carried out under zoo conditions.
The positions of males and females during the emission of urine are practically
indistinguishable. In cases where rocks and trees are being marked, the
animals do not squat down--"walking up to the object, they first sniff it,
then they turn backwards toward it, raising their tail vertically above the
site to be marked (Fig. 20). A stream of urine, which
is often atomized, is ejected directly backward for a distance of 20-30 cm
or less. In nature, such urine marks are detected by the more or less abrupt
turns made by the tigers toward individual tree trunks, tree stumps, or rocks
(Fig. 21). Sometimes "dirty footprints" are left at
such a site, but only occasionally are there individual drops of urine that
fall onto the snow. Urine markings that are left by tigers on tree trunks
and rocks along the tigers' paths without the tiger changing the direction
in which it is walking can be easily missed by the observer" (Matyushkin
& Yudakov 1974, p. 16). Urine markings on vertical surfaces have been
found at heights of 60-125 cm, most often at about a meter from the level
where the prints of the tiger's paws were left by the animal at the base
of the marked object (Fig. 22). In entries made in
field notebooks, these turnings of tigers toward marked objects are briefly
designated as "approaches".
The other most common form of marking behavior of tigers is "scraping",
as a result of which there form easily noticeable streaks of snow or soil
that have been dug up by the tiger [so-called "scrapes" or "rakes" (of the
tigers claws), Fig. 23]. They are considered "clear
evidence that the footprints belong to a male" (Yudakov 1974, p. 355).
Feces--in large or small amounts (Fig. 24), up to
several drops of semi-fluid feces ("punctate excrement")--are left behind
on scrapes in various situations, as well as the spots that are evidence
of the copiousness of urine emissions, or only the aggregation of a splatter
of urine that had penetrated the snow. Not infrequently, scrapes are observed
that seem "empty" upon a superficial examination. However, it is necessary
to take into consideration that the small urine marks on them are difficult
or impossible to discover, since needles or branches of conifers, dry leaves,
other kinds of plant debris, etc. lie above them.
Sometimes, when approaching trees or rocks, tigers stretch upward onto
them, leaning the front part of their bodies on such objects and scratching
the trunks. The traces that remain after this on the vertical surfaces
can also fulfill a marking
function.
We observed rubbing in 11 (2.5%) and scratches in 16 (3.7%) of all "approaches".
The upper levels of the scratch marks, which were left on tree trunks by
different individuals, were found at heights ranging from 210 to 290 cm.
Possibly, the behavior of the animals when the tigers break seedlings of
coniferous trees into pieces or gnaw around branches, and then lie down on
them or roll on them also has marking significance. We observed this phenomenon
14 times.
In order to give an idea of the frequency of different forms of marking behavior during the journeys of tigers and of the place that they occupy in the usual sequence of behavioral acts, we present two specific examples. On February 27, 1972, we tracked the footprints of the "Emperor" male tiger, which had gone along a spur of the right bank of Bogdanov Stream in the direction of the valley of the Gornaya River. The animal's path ran along a flattened, smooth spur, which had smooth contours, gradually descending into the valley. The forest at this place is Korean Pine forest with an admixture of other dark coniferous species.
Figure 19. A shelter (or lair) used by tigers under a rock overhang.
Figure 20. A tigress places a urine mark on the wall of her cage (Moscow
Zoo).
Photo: E. N. Matyushkin.
Figure 21. An approach by the "Emperor" male tiger to an object, the "stump", that he constantly marked.
At a distance of 220 m from the beginning of the sector of the route
selected for description, the tiger walked beneath a dead and withered tree
that was leaning to one side and "marked" it. After going 880 m farther,
he had lain down for a short while. At a distance of 3 m from his laying
place, he made a scrape and left a small amount of excrement on it. Upon
going another 70 m, the tiger rolled around in the snow for a long time.
The subsequent 370 m sector of our tracking terminated at a site where the
tiger had made a urine mark on a stone, which had a shape reminiscent of
that of a mushroom. Immediately beyond the rock, there was an "empty" scrape.
Only 330 m of the tiger's path separated this site from the next laying
place, but, at a distance of 110 m beyond the laying place, we discovered
a scrape and a urine mark, again on a dead, withered tree that was leaning
to one side. After going 140 m farther, the tiger had lain down for a while
under a tree, which he later scratched. After going another 110 m, the tiger
had lain down under a Korean pine tree, and, at a distance of 70 m from that
site, he "marked" a sloping oak tree. After going another 70 m. along the
tiger's tracks, we found a new laying place. After going 65 m farther, there
was a place where the tiger sat down; slightly farther on, he had lain down.
After going 400 m farther, the animal reached the valley of the Gornaya
River, and, soon, after walking another 160 m, the tiger reached the road
running along this river. The total length of the sector of his path that
we have examined is about 3 km.
On February 28 of the same year, we tracked the "Empress" tigress near
the site that we have just described. At a distance of 70 m from the beginning
of the path that we had selected in the nature of an example, she had lain
down underneath a birch tree for a short time. Then she approached a neighboring
tree and left a urine mark on the trunk. After going 145 m farther, the
tigress trampled around a dead, withered tree. After going another 70 m.,
she had lain down for a while under a Korean pine tree and scratched it.
From this point onward, she changed the direction of her movements to the
opposite direction. After going a distance of 330 m during her descent from
a ridge, we observed a characteristic marking approach by the tigress to
a Korean pine tree. The subsequent 145 m of tracking led to a laying place
used briefly by the tigress. After going a distance of 400 m, the tigress
made an "approach" to a dead, withered tree, and, after proceeding for another
590 m, there were footprints indicating "trampling or kneading by the tigress'
feet" near a dead, withered tree that was leaning to one side.
Figure 22. A urine mark at the base of a shrub that was made by the
"Emperor" male tiger at the time of the rut.
Figure 23. A scrape left by a male tiger on the terrace of a stream.
After the tigress had walked for 1500 m downward along a slope into the
valley of Kamennyi Stream, she traversed the channel of a stream after going
a further 370 m. Then, after going another 250 m, the tigress emerged onto
a road. When going along the road, the tigress at first moved upward (relative
to the elevational gradient of the valley), but she covered a distance of
only 30 m in this direction, and, having left a urine mark, she abruptly turned
around. Now proceeding downward along the road, she approached a tree on
the verge of the road after going another 140 m in a steeply arced path.
Then she kneaded with her paws around a dead fallen tree. Precisely here,
turning to her right, the tigress left the road and headed along a track
going toward a mountain. She had lain down three times over a distance of
845 m while making her ascent. From the last laying place, which was near
the crest of a spur, the tigress made a 20-meter approach to a large birch
tree, under which she kneaded with her feet, making a circle. Here the tigress
abruptly altered her "course", heading on a path that made an acute angle
with her line of movement prior to this. Now, she headed upward along the
crest of the spur: she lay down after going 130 m; once again she had lain
down after going another 55 m. At a distance of 25 m from there, the series
of footprints described a sharp salient point
of a curve under a birch tree. Walking for another 350 m, the tigress turned
underneath sloping dead and withered tree. Only 180 m separated this site
from her next laying place. The total path length of the animal in the present
example was approximately 5.5 km.
From the individual examples cited above, it is already possible to note
important features of the marking activity of tigers. First, marking behavior
belongs, together with the laying places used briefly by tigers, among the
most frequent behavioral
acts recorded during our tracking. The distance between neighboring marks
is frequently measured in several hundreds or even several tens of meters
(Fig. 25). In the case of males, the scrapes and markings
on vertical surfaces alternate without
any sort of rule governing their sequence. Second, it is noteworthy that
distinct markings or combinations of markings (Fig. 26)
are almost always found in those locations where the animals abruptly change
the direction of their path, where they encounter either the tracks of other
tigers or the tracks of a human (and the evidence of that person's activity).
Third, tigers mainly select for marking those objects that in some way stand
out from the surrounding background. Thus, trees of large diameter are clearly
preferred over thinner trunks, and leaning trees are marked more frequently
than upright trees. The tigers are attracted to dead,
withered trees and to trees and rocks of unusual shapes. It is also necessary
to add sawed off logs that protrude from the verges of roads and paths (Fig. 20), tree trunks, and fallen trees ("torchiny," trees
that stand out from the background) to
the list of favorite objects for marking.
Figure 24. A scrape (with excrement) made a male tiger.
Marking Activity of Male Tigers on Journeys With Varying Degrees of
Prominence of Visual Landmarks.
| Section of the Route (in parentheses,
the distance of tiger prints tracked, in km)
|
Urine Marks on Vertical Surfaces (tree trunks, rocks, etc.) |
Urine Marks on the Surface of the Snow (without scrapes) |
Excrement (without scrapes) |
|||
|
|
Total |
Per 10 km |
Total |
Per 10 km |
Total |
Per 10 km |
| Along forest massifs (stands)
and in river valleys without clear visual landmarks (434 km) |
109 | 2.5 | 21 | 0.48 | 17 | 0.39 |
| Along crests of mountain spurs
(72.4 km) |
69 | 9.5 | 1 | 0.14 | 7 | 0.97 |
| Along terraces and foothills
of slopes (43 km) |
45 | 10.5 | 2 | 0.46 | 7 | 1.5 |
| Along roads (115 km) | 96 | 8.35 | 2 | 0.17 | 7 | 0.6 |
| Total (664.4 km) | 319 | 4.80 | 26 | 0.39 | 38 | 0.57 |
| --cont | Scrapes: | |||||||||
| "Empty" | With a small amount of urine | With large spots of urine | With excrement | Total | ||||||
| Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
|
|
Along forest massifs |
33 | 0. 76 |
42 | 0.97
|
4 | 0.09
|
12 | 0.28
|
91 | 2.1 |
|
Along crests of mountain spurs |
31 | 4.28 | 32 | 4.42 | 3 | 0.41 | 11 | 1.52 | 77 | 10.6 |
|
Along terraces and foothills of slopes |
17 | 3.95 | 13 | 3.02 | - | -
|
2 | 0.46 | 32 | 7.44 |
| Along roads |
27 | 2.35 | 40 | 3.48 | 2 | 0.17 | 19 | 1.65 | 88 | 7.65 |
| Total |
108 | 1.6 | 127 | 1.9 | 9 | 0.13 | 44 | 0.66 | 288 | 4.33 |
Marking Activity of Tigresses on Journeys With Varying Degrees of
Prominence of Visual Landmarks.
| Sections of the Route (in parentheses, the distance of tiger prints tracked, in km) |
Urine Marks on Vertical Surfaces (tree trunks, rocks, etc.) |
Urine Marks on the Surface of the Snow (without scrapes) |
Excrement (without scrapes) |
|||
|
|
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
| Along forest massifs (stands)
and in river valleys without clear visual landmarks (304.6 km) |
69 | 2.26 | 21 | 0.7 | 36 | 1.2 |
| Along crests of mountain spurs (29 km) | 22 | 7.6 | 3 | 1.0 | 8 | 2.7 |
| Along terraces and foothills of slopes (25.4 km) | 7 | 2.7 | 3 | 1.2 | 1 | 0.4 |
| Along roads (17.5 km) | 11 | 6.28 | 1 | 0.6 | 1 | 0.6 |
| Total | 109 | 2.90 | 28 | 0.7 | 46 | 1.2 |
| --cont. | Scrapes: | |||||||||
| "Empty" | With a Small Amount of Urine |
With an Abundance
of Urine Spots |
With Excrement | Total |
||||||
| Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
|
|
Along forest massifs |
- | - |
|
- |
|
- |
|
- | ||
|
Along crests of mountain spurs |
- | - | - | - | - | |||||
|
Along terraces and foothills of slopes |
- | - | - | | - | - | ||||
| Along roads |
1 | 0.6 | - | - | - | 1 | 0.6 | |||
| Total |
1 | 0.03 | - | - | - | 1 | 0.03 | |||
Selectivity in the arrangement of markings is still more graphically reflected
in the estimated totals relative to the total aggregate of the data (Table 12). Among the objects marked by tigers, upright
trees are almost 1.5 times more numerous than leaning trees. However, for
the long-term study area as a whole, typically the proportion of upright
trees exceeds by 5-9 times the percentage of leaning trees (we have in mind
trees that are at least 25-30 cm in diameter). Therefore, the tigers marked
the latter kind of tree (i.e., leaning trees) 3-6 times more readily. The
comparison of the proportion of dead trees among those trees marked by tigers
with the percentage of dead trees in timber stands allows us to conclude
that the animals prefer dead, withered trees for marking.
Associations Between Markings on Vertical Surfaces (n=379) and Various
Objects.
| Object Marked |
Frequency | |
| | Absolute frequency |
% |
| Living Trees: | 260 | 68.60 |
| Live trees standing upright | 154 | 40.60 |
| Live trees leaning to one side | 106 | 28.00 |
| Dead Standing Trees: | 37 | 9.75 |
| Dead trees standing upright | 18 | 4.75 |
| Dead trees leaning to one side | 19 | 5.00 |
| Sawed off logs, dead fallen wood, stumps | 64 | 16.90 |
| Rocks, Stones | 18 | 4.75 |
Specific tree species are also preferentially marked. Most often, tigers leave marks on birch trees (Betula sp.), in particular silver birch (B. costata) [these trees do not dominate among the standing timber and contrast with those that surround them]. Out of 112 markings for which the species of tree was indicated, 63 were on birch (56.2%); 16 (14.3%) were left on linden trees (Tilia spp.), and 14 (12.5%) were on Korean pine trees. The remaining 19 markings (17%) were associated in approximately equal numbers with the other tree species. It is remarkable that tigers usually do not walk past the yew trees (Taxus spp.) that are sometimes
encountered along their routes.
Along the paths traveled by tigers, there exist objects that are invariably
marked, which the tigers almost never leave without giving them their attention,
whenever the tigers appear in the vicinity of such objects (cf., Fig. 20). In exactly the same way, the animals can make
scrapes (Fig. 25) in one and the same place when repeating
their routes. A distinctive microbiotopic association characterizes the
latter type of mark (as is also true of urine marks on vertical surfaces).
For example, tigers often leave scrapes where a thin undergrowth of coniferous
species is present, the animals sometimes breaking up these seedlings while
making their scrapes (Fig. 26). Often the tigers display
marking activity in the direct vicinity of their laying places (Fig. 27).
Not only different objects and locations in the individual territories
of the animals, but also different sections of their routes have different
probabilities of being marked (even if the number of sites suitable for marking
is the same everywhere). Observations showed that this inequality depends
not only upon the composition and density of the timber stands, but also
upon whether the tiger's path runs along a valley or a mountain slope. A
clear connection was discovered between the frequencies with which tigers
applied marks and the degree of prominence of visual landmarks along the
tigers' routes (cf., Tables 10 & 11). Since it is precisely such landmarks that serve
as a prerequisite for the selection by the animal of a completely determined,
but not an arbitrary route, the conclusion was drawn that the animals above
all mark "sites near which other tigers are most likely to pass" (Yudakov
1974, p. 355). Thus, in the case of males, the frequency of encounter of
scrapes during the tiger's movement along the crests of mountain spurs is
five times greater than during travels in homogeneous forest massifs (dense
stands of tall trees). A similar pattern is revealed for "approaches".
In males, the maximum frequency of "approaches" was recorded for sectors
of the tigers' routes that ran along river terraces and foothills of slopes
as well as along crests of mountain spurs and roads. The same tendencies
were traced in the changes in marking activity of tigresses: they most intensively
marked their routes along mountain spurs and along roads. Under conditions
where homogeneous forest massifs are present, the number of "approaches"
per unit of distance (relative to the maximum number of "approaches") falls
by approximately a factor of three.
Figure 25. Scrapes made by a male tiger, with one scrape following
immediately after the previous one. At the center are two scrapes left at
one site at different times.
Figure 26. A combination of different forms of marking: a scrape made
by a male tiger with his subsequent approach to the trunk of a Korean pine
tree (a urine mark on the vertical surface).
Figure 27. A scrape made by a male tiger near his laying place (which is melting away (melting on the outside)).
Together with features that are similar for the marking behavior of
females and males, we observed essential differences. Males mark their routes
more intensively: the frequency of "approaches" is 1.5 times higher in males.
Only on the sectors of tigers' routes along mountain slopes and river valleys
did we obtain similar indices for animals of both sexes. But even in the
latter case, the equal frequency found for the two sexes is true only of
urine markings on vertical surfaces. With regard to scrapes, the differences
among the sexes contrast greatly. If, in the case of males, scrapes are
encountered almost as frequently as urine marks on trees, stumps, and rocks,
then for tigresses this form of marking was observed to be the exception.
Speaking more accurately, a scrape that probably belonged to a female was
observed only once during the entire period of our study. We did not observe
that females broke up or gnawed around the stems of seedlings of coniferous
trees nor did they rub themselves on trees, although the latter behavior
could have been completely missed by us. Scratches on trees that were left
by females were observed to occur eight times less frequently (than in males).
Marking Activity of Tigers During the Rut and Outside the Mating
Period.
| Sex and Physiological State of the Individual |
Urine Marks on Vertical Surfaces |
Urine Marks on the Surface of the Snow |
Excrement
|
|||
|
|
Total |
Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
| The "Empress" Tigress outside
the period of sexual activity (159 km of tracking) (1) |
32 | 2.01 | 3 | 0.2 | 9 | 0.6 |
| The "Empress" Tigress during
estrus (119 km of tracking) (2) |
67 | 5.6 | 8 | 0.7 | 16 | 1.3 |
| The "Emperor" male outside the period of sexual activity (319 km of tracking) (3) | 85 | 2.66 | 16 | 0.5 | 14 | 0.4 |
| The "Emperor" male having joined
the female during her estrus (141 km of tracking) (4) |
66 | 4.68 | 3 | 0.2 | 16 | 1.13 |
| --cont. | Scrapes: |
|||||||||
| "Empty" scrapes |
With a small amount of urine | With abundant urine spots | With excrement | Total | ||||||
| Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
|
|
(1) |
- | |
|
|
|
|
|
- | ||
|
(2) |
1 | 0.08 | - | - | - | 1 | 0.08 | |||
|
(3) |
41 | 1.3 | 17 | 0.5 | 7 | 0.2
|
6 | 0.19 | 71 | 2.2 |
| ( 4) |
24 | 1.7 | 49 | 3.47 | - | 27 | 1.91 | 100 | 7.09 | |
Marking Activity of Male Tigers in Different Parts of Their Territories
(based on the tracking of the "Emperor" male).
| Location of the Observation
Site Relative to the Entire Area of the Territory |
Urine Marks on Vertical Surfaces | Urine Marks on the Surface of the Snow | Excrement |
|
|||
| Total |
Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
||
| "Core" of the Territory (319 km of tracking) | 85 | 2.66 | 16 | 0.5 | 14 | 0.4 |
|
| Periphery of the Territory (33 km) | 25 | 7.57 | | | | |
|
|
|
Scrapes: |
|||||||||
| "Empty" scrapes | With a small amount of urine | With an abundance of urine spots | With Excrement | Total | ||||||
| Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
Total | Per 10 km |
|
|
|
41 | 1.3 | 17 | 0.5 | 7 | 0.2 | 6 | 0.19 | 71 | 2.22 |
|
|
17 | 5.1 | 6 | 1.8 | | | 1 | 0.3 | 24 | 7.27 |
*Data for the period during the rut are not included in the Table.
The marking activity of both tigresses and male tigers increases substantially
during the mating period (Table 13). "Approaches" began
to be encountered approximately twice as frequently along the tracks of females
that were in estrus. The intensity of marking of vertical surfaces by male
tigers that are searching out females or that have already joined up with
them during estrus changes about as much. The number of scrapes per unit
of distance during this time increases still more abruptly, by approximately
a factor of three.
In the case of females with cubs, marking activity is, on the contrary, noticeably
less frequent.
A question of great interest is: how does the intensity of marking change
from the center of the territory to its periphery? Answering this question
is very complicated, since we have not yet succeeded in clearly delimiting
the center and
periphery of the territories. Moreover, even the data from regular tracking
over the course of three winter seasons is insufficient to solve such a problem.
Nevertheless, we attempted to compare the results of observations on the
"Emperor" tiger in those sites where he walked most frequently with those
places where he rarely went, but where a zone of contact in the valleys and
slopes of Petrov and Zvuchnyi (Nikolaeva) Streams clearly existed between
the territory of this animal and that of his neighbor, the "Powerful" male
tiger. Let us remember that it was exactly here that the nomadic male, the
"Lazy" tiger, perished in a fight with
the "Powerful" tiger. A comparison shows (Table 14)
that the intensity of marking of his territory by the "Emperor" tiger was
approximately three times higher in the zone of contact between the territories
of the two males. Increased marking activity
was also recorded here in the case of the "Powerful" tiger up until his encounter
with the "Lazy" tiger. The differences expressed relate equally to urine
marks on vertical surfaces and to scrapes.
With respect to the case under discussion, we have possibly come across
the manifestation of a general principle, but additional observations are
needed for the support of such a conclusion. However, even if the periphery
of the territory is actually marked more actively, there is no basis for
speaking of a preference existing for marking the borders of the individual
territory as such (i.e., the perimeter of the territory). We have
discussed earlier the absence of clear-cut boundaries as such. Routes that
are frequently repeated by the tigers are marked to a greater degree, both
on the periphery of the territory as well as in its "core".
The degree of prominence of visual landmarks along the routes and the probability
of contact between a given individual and other tigers (which is connected
with such landmarks) are factors of the first rank that influence the intensity
of marking. The
statements expressed above are supported, in particular, by the fact that
the nomadic male, the "Lazy" tiger, left marks along his routes not only
not less frequently but even significantly more often than did the "Emperor"
tiger which constantly lived here (for "approaches": 7.2 vs. 3.6 per 10 km;
for scrapes: 5.5 vs. 3.9 over the same distance). We stress once again that
the "Lazy" tiger utilized roads for his travels to a still greater degree
than did other individuals.
On the whole, according to the data that we possess, marking behavior in
the Amur tiger can be considered as an important mechanism allowing: (1)
the maintenance of stable connections between each individual and a defined
territory, (2) setting internal borders of the territories of animals in
relation to a an established type of spatial structure of the population,
and (3) the preservation of constant contacts between individuals that are
living in the same vicinity.