F:\ALCES\Supp2\PAGEMA~1\Rus15s.


ALCES SUPPL. 2, 2002 KUZNETSOV – MOOSE AND FOREST PROBLEMS IN RUSSIA

65

MOOSE AND FOREST PROBLEMS IN RUSSIA

German V. Kuznetsov

Institute of Evolutionary Animal Morphology and Ecology, Russian Academy of Science, 117071,

Moscow, Russia

ABSTRACT: This article presents an analysis of the moose–forest relationship in Russia charac-

terized by utilization of land by humans and its consequences for moose and the forest.  It provides

a general overview of the research approaches regarding Russia’s damaged forests by moose.  In

the early 1950s, the moose population increased sharply, primarily due to enlargement of the cutover

area and the ensuing increased forage resource.  Devastation to pine and oak are emphasized amid

a backdrop of damage to silviculture that cost millions of rubles.  Other northern countries were

undergoing similar destruction by moose to their forests.  Three main research approaches are

distinguished: determination of the damage by moose to stands, estimation of the effects of moose

on the structure of forest phytocenoses, and the effects of moose on the productivity of particular

plant species and forest phytocenoses.  This well–documented article correlates various moose

population densities with specific effects on different ecosystems and emphasizes the fact that

trophic activity of moose is one of several factors affecting the structure and succession of forest

phytocenoses of various natural zones.

ALCES SUPPLEMENT 2: 65-70 (2002)

Key words: biocenosis, ecology, forest, moose, phytocenoses, population density, productivity,

succession, silviculture, trophic activity

In the USSR, a moose and forest prob-

lem originated in the early 1950s, when the

moose population increased sharply, prima-

rily due to enlargement of the cut over area

and, subsequently, the increased forage re-

source.  In this situation the moose began to

detrimentally affect young forest growth

and regrowth opportunity, especially those

of pine and oak.  The damage inflicted by

moose on silviculture was estimated at mil-

lions of rubles.  A negative effect of the

moose on the forest was also recorded in

other countries, such as Sweden and Nor-

way (Yurgenson 1979, Filonov 1983).

Analysis of the studies conducted in the

USSR on the moose and forest problem

shows 3 main approaches: (1) determina-

tion (%) of the degree of damage by moose

to forest stands; (2) evaluation of the effect

of moose on the structure of forest

phytocenoses and succession processes;

and (3) investigation of the effect of moose

on the productivity of particular plant spe-

cies and forest phytocenoses.  These 3 main

approaches to a certain extent reflect the

history of the development of research into

estimation of the functional role of moose in

forest biocenoses.  In most studies of the

first approach, the damage estimate of moose

on the forest was based on the criterion of

the level of damage to the shoots of trees

(%).  However, on the basis of such data,

the effect of moose on the productivity of

forest phytocenoses and related compo-

nents cannot be estimated.  Nevertheless,

an examination of studies on this subject is

valuable to address the large body of evi-

dence already collected from these data.

An inverse relationship between the

area of pine plantations and the rate of their

damage by moose was established; i.e., the

less area of pine plantations on cutovers



MOOSE AND FOREST PROBLEMS IN RUSSIA – KUZNETSOV ALCES SUPPL. 2, 2002

66

being overgrown, the greater they are af-

fected by moose (Dinesman 1961).  It fol-

lows that plantations should have a great

area to resist the trophic pressure of moose.

These results were supported by further

studies.  In fact, a number of authors

(Kozlovsky 1960, Kaletskaya and Kudinov

1987) revealed that a ratio of 20–30 ha of

forage grounds (forest plantations) per

moose reduces the detrimental role of moose

to a minimum and, conversely, a ratio of <10

ha per moose sharply increases the damage

to plantations by moose.  To some degree

this index indicates the expected effect of

moose on forest plantations.  For instance,

in the Nizhny Novgorod Region, the area of

young pine per moose is 112 ha, while in the

Novgorod Region, it is only 4.3 ha

(Chervonnyi 1975).  Naturally, in the

Novgorod Regions the effect of moose on

forest plantations is more substantial.

Judging from the fact that dense planta-

tions of pine are damaged by moose to a

much lesser extent, Borodin (1959b) pro-

posed a biological approach to pine protec-

tion from moose through an increase in the

density of its plantations.

Numerous authors (Borodin 1959a,

Dinesman 1961, Kheruvimov 1969, Padaiga

1980, Smirnov 1987) have demonstrated

that intensive trophic activity of moose de-

lays the growth of pine, oak, ash, aspen,

birch, fir, and other species, which is re-

flected in the quantity of timber.  Subse-

quently, a number of researchers revealed

that the intensity of forest damage by moose

is a function of their population density.  In

f a c t ,  i n  s m a l l  f o r e s t  m a s s i f s  o f  t h e

Tsentralno–Chernozemny Reserve, the

trophic pressure of moose is fairly heavy

(Gusev 1988).  It is understandable that the

degree of moose effect on forest planta-

tions largely depends on the absence, in

some cases, of a clear cut relationship be-

tween the population density of moose and

plantation productivity over large areas

(Kuznetsov 1980).

A large body of evidence was obtained

on the development of stands under the

effect of moose in reserves of the USSR;

Prioksko–Terrasny, Oksky, Darvinsky, etc.

( K a l e t s k a y a  1 9 5 9 ,  K o z l o v s k y  1 9 6 0 ,

Timofeeva 1974, Chervonnyi 1975, Dunin

1975, Zablotskaya and Zablotskaya 2002).

The effect of moose is one of a series of

factors affecting the health of the stand.  In

mixed plantations, it is important that such

factors as growth conditions, the ratio of

different trees, and the distribution of avail-

able forage should be taken into account.

Necessary cutovers can promote the for-

mation of high–quality timber.  The recent

discussion on the moose–forest problem in

t h e  j o u r n a l  O k h o t a  i  O k h o t n i c h y e

Khozyaistvo (Hunting and Game Manage-

ment) demonstrated that there are many

problems yet to be solved on rational use of

moose and the forest (Pavlinov 1983, Dunin

1984, Perovsky 1984).

A second direction of research is that it

is known that moose promote replacement

of the main species of regrowth and

underbrush and the specificity of the effect

of moose is associated with the features of

succession in a particular region.  For in-

stance, on Valdai, as a result of the effect of

moose, there occurred gradual replacement

of pine by deciduous trees and spruce.

Since spruce forests form under natural

succession, there are grounds to believe

that the moose promoted acceleration of

this process.  Under conditions of the south-

ern taiga, moose under different population

density exert a dissimilar effect on the de-

velopment of forest stands.  When moose

numbers are low, spruce–birch stands de-

velop, but when moose numbers are high,

primary spruce stands recover.  An exces-

sive population of moose degrades stands

(Smirnov 1987, Abaturov and Smirnov 2002).

In another region in Tulskie Zaseki, accord-

ing to our data, moose damaging oak trees



ALCES SUPPL. 2, 2002 KUZNETSOV – MOOSE AND FOREST PROBLEMS IN RUSSIA

67

promote acceleration of the growth of ash,

in particular the linden and filbert.  There,

the moose acts as a factor promoting the

development of shady linden forests.  Thus,

the trophic activity of moose is one of the

factors affecting the structure and succes-

sion of forest phytocenoses of various natu-

ral zones.

Another approach to the moose–forest

problem is associated with the investigation

of the role of moose in the productivity of

particular species of arboreous and herba-

ceous plants and phytocenosis.  The main

data were obtained through comparison of

the state and productivity of plants under

the effect of moose and in isolation from

them.  Under conditions of forest–steppe,

moose affect the growth and development

of broad–leaved forests and decrease their

productivity (Zlotin and Khodashova 1974,

Gusev 1989).  In Tulskie Zaseki, oak and

ash respond differently to the effect of

moose.  Under conditions of a constant

effect of moose, the oak reduces its produc-

tivity even under a small level of removal

(13%).  Conversely, the ash is more resist-

ant to removal of its phytomass by moose,

which appears to be due to the biological

properties of this species; on average, the

annual increment of the leaf phytomass

exceeds tenfold the respective increment of

shoot phytomass.  The shoots of ash are

protected by the mass of foliage, and, hence,

the moose utilizes them to a less extent.

In Valdai under conditions of the south-

ern taiga, the annual increment in the Scots

pine in isolation was twice as high as in the

pines accessible to moose; the admissible

removal of annual shoots under which pro-

ductivity is not reduced is close to 50%.

It should be noted that removed phytomass

of the pine is used by the moose fairly

effectively and forage remains about 5%.

It should also be emphasized that the state

and productivity of pines is a function of not

only the effect of moose but also of a set of

other factors.  For instance, the productivity

of pines in an elevated plot is twice as high

as in a lowland bogged area, despite the fact

that the trophic pressure of moose in the

former case is tenfold higher (Kuznetsov

1980).

The effect of moose on spruce is deter-

mined to a great extent by penetration of

insect pests through injured parts of the

stalk and infection by timber rots, which

results in disintegration of the spruce layer

of the stand rather than by removal of

regrowth (Smirnov 1987).  Willow and

mountain ash can sustain 70–80% of re-

moval of annual increment over many years,

the productivity being maintained at a rela-

tively constant level.  The mountain ash is

more resistant to phytomass removal by

moose than birch and willow; favorite for-

age of moose is still moose resistant

(Kuznetsov 1980, Chernyavsky and Dubinin

1989).

A cycle of observations over the natural

response of arboreous plants to removal of

their increment by moose is supplemented

by data on experimental removal of

phytomass in the pine and oak, and aspen

(Smirnov 1987).  It was demonstrated that

the productivity of the pine depends not only

on the amount of removal of the increment

phytomass but also on the method by which

it is removed; i.e., removal of individual

shoots does not bring about as sharp a

reduction in productivity as does removal of

the same amount of phytomass from each

shoot.  The moose normally browse only a

part, about one third, of the shoots; it chooses

the most judicious method of using the in-

crement.  Also, as shown by defoliation and

cutting the shoots, aspen sensitivity to re-

moval of shoots can serve as a good indica-

tor of the state of forage resources of

moose (Smirnov 1987).  In general, we can

conclude that deciduous trees, especially

those that have root shoots, sustain greater

trophic pressure by moose than coniferous



MOOSE AND FOREST PROBLEMS IN RUSSIA – KUZNETSOV ALCES SUPPL. 2, 2002

68

and those without shoots.

How moose actively affect the produc-

tivity of forest phytocenosis is not fully

understood; however, the importance of

this line of research is beyond doubt.  We

have demonstrated in Valdai that such char-

acteristics as annual increment, the long–

term reserve of phytomass of herbaceous

vegetation and shrubs, and the annual incre-

ment of heath under the effect of moose

exceed the respective parameters under

conditions of isolation.  The increase in

productivity of the pine in an open area (in

a “range”) appears to be compensated for

by an increase in the total increment of

herbaceous plants and shrubs at the level of

phytocenosis.  However, to get the total

balance of phytocenosis productivity, a

number of other parameters are needed

such as increment of the stalk loss due to

trampling, etc.  But still there are grounds to

believe that moose, affecting the value of

annual increment in some individual plant

species, can to a considerable degree re-

duce the productivity of entire phytocenoses

(Kuznetsov 1980).

Thus, the line of research in question

makes it possible to estimate the activity of

moose as a component of forest ecosys-

tems.  The ecosystem approach regards the

activity of moose as the same damage in-

flicted on the forest, since primary produc-

tion will be transformed into secondary and

the very notion of “damage” does not exist

in the biological sense.  Conversely, the

silvicultural approach presupposes obtain-

ing high–quality timber and, hence, the moose

and forest problem becomes realistic.  In

this connection, it is not by chance that the

efforts of both zoologists and silviculturalists

are aimed at investigating methods of forest

protection from moose.  The most harmless

and accessible are the biotechnical meth-

ods, such as supplemental feeding, planting

of fast–growing shrubs, and shoot cutting,

etc.  However, when moose numbers are

high, these methods yield no positive effect.

Presumably, under these conditions it would

be expedient to use such a powerful method

of moose population control as removal.

But in this country, only 10–12% of the

moose population is removed, which is ob-

viously insufficient.  It will be remembered

that in Sweden they removed 40–50% of

the population (Dezhkin 1983).  Of interest

are the approaches to the investigation of

the intrapopulational structure and its role in

the regulation of moose numbers (Baskin

1984).

Mechanical methods, the exclosure of

commercially important arboreous species,

can be the most effective, but they are

associated with great economic expense

because exclosures must be efficient to

achieve the desired outcomes.  The optimal

methods for protection of forest from moose

depend on particular conditions in specific

regions, and their application should be inte-

grated.  In recent years chemical methods

of forest protection from moose have been

developed (Martynov 1980).  The applica-

tion of repellents can in some regions be

very promising, but, unfortunately, there are

no data available on the genetic control of

their application.  Thus, through the efforts

of scientists, the set of methods of forest

protection from moose increases, but their

implementation is lagging behind.  It can be

hoped that increasing interest in the prob-

lems of conservation of the natural environ-

ment will stimulate the solutions for these

practical problems in the field of moose

ecology.

REFERENCES

ABATUROV, B. D., and K. A. SMIRNOV.  2002.

Effects of moose population density on

development of forest stands in central

European Russia.  Alces Supplement

2:1-5.

BASKIN, L. M.  1984.  Moose. Pages 45–73

in V. E. Sokolov, editor.  The wildlife of



ALCES SUPPL. 2, 2002 KUZNETSOV – MOOSE AND FOREST PROBLEMS IN RUSSIA

69

the southern taiga.  Nauka, Moscow,

Russia.  (In Russian).

BORODIN, L. P.  1959a.  On the problem of

moose role in silviculture.  Proceedings

of the Institute of Silviculture, USSR

Academy of Sciences 3:102–110.  (In

Russian).

.  1959b.  Increase in the density of

pine plantations as a biological method

of pine protection from the moose.

Proceedings of the Institute of Silvicul-

ture, USSR Academy of Sciences

13:124–126.  (In Russian).

CHERNYAVSKY, P. B., and V. N. DUBININ.

1989.  Moose in northeastern Siberia.

Nauka, Moscow, Russia.  (In Russian).

CHERVONNYI, V. V.  1975.  Winter forage

resources and feeding of moose in the

European RSFSR.  Proceedings of the

Oksky State Reserve 11:321–389.  (In

Russian).

DEZHKIN, V. V. 1983.  Hunting and game

management of the World.  Lesnaya

Promyshlennost, Moscow, Russia.  (In

Russian).

DINESMAN, L. G.  1961.  The effect of wild

mammals on the development of forest

stands.  USSR Academy of Sciences,

Moscow, Russia.  (In Russian).

DUNIN, V. F.  1975.  The effect of moose on

forest phytocenoses of Byelorussian

LakeArea. Pages 167–168 in Ungu-
lates of the USSR fauna.  Nauka, Mos-

cow, Russia.  (In Russian).

.  1984.  Both forest and moose.

Okhota i Okhotnichye Khozyaistvo 2:8–

9.  (In Russian).

FILONOV, K. P. 1983.  Moose.  Lesnaya

Promyshlennost, Moscow, Russia.  (In

Russian).

GUSEV, A. A.  1988.  Admissible density of

the population of wild ungulates and

experience of its maintenance in the

Tsentralno–Chernozemny Reserve.

Pages 114–128 in Population studies of
animals in reserves.  Nauka, Moscow,

Russia.  (In Russian).

.  1989.  Animals in reserved terri-

tories.  Tsentralnoye Chernozemnoye,

Moscow, Russia.  (In Russian).

KALETSKAYA, M. A., and A. A. KUDINOV.

1987.  Development of pine plantations

of dense young stands damaged by

moose.  Pages 189–215 in A. G.
Bannikov, editor.  Biology and harvest

of moose.  Volume 3.  Moscow, Russia.

(In Russian).

KALETSKAYA, M. L. 1959.  Damage by

moose to pine seedlings in the Darwin

Animal Preserve.  Proceedings of the

Institute of Silviculture, Russian Acad-

emy of Sciences 13:63–69.  (In Rus-

sian).

KHERUVIMOV, V. D. 1969.  Moose (Com-

parative Studies as Exemplified by the

Tambov Population).  Tsentralno–

Chernozyomnoye, Moscow, Russia.  (In

Russian).

KOZLOVSKY, A. A. 1960.  Protection of

f o r e s t s  f r o m  d a m a g e  b y  m o o s e .

VNILM, Moscow, Russia.  (In Rus-

sian).

KUZNETSOV, G. V. 1980.  The role of

ungulates in forest ecology (some

insights and perspectives from re-

search).  Pages 88–100 in Herbivores
in vegetative communities.  Nauka,

Moscow, Russia.  (In Russian).

MARTYNOV, E. N. 1980.  The effect of

chemical treatment of forest on the

b i r d s  a n d  m a m m a l s .   L e s n a y a

Promyshlennost, Moscow, Russia.  (In

Russian).

PADAIGA, V. I.  1980. Estimate of the dam-

age inflicted on silviculture by Cervidae.

M e t h o d o l o g i c a l  r e c o m m e n d a t i o n .

Kaunas, Lithuania. (In Russian).

PAVLINOV, N. 1983.  Moose through the

eyes of a silviculturist.  Okhota i

Okhotnichye Khozyaistvo 7:10–11.  (In

Russian).

PEROVSKY, M. D. 1984.  Moose and forest:



MOOSE AND FOREST PROBLEMS IN RUSSIA – KUZNETSOV ALCES SUPPL. 2, 2002

70

aspects of equlibrium.  Okhota i

Okhotnichye Khozyaistvo 2:9–10.  (In

Russian).

SMIRNOV, K. A. 1987.  The role of moose in

biocenoses of the Southern Taiga.

Nauka, Moscow, Russia.  (In Russian).

TIMOFEEVA, E. K. 1974.  Moose: ecology,

distribution, economic importance.  Len-

ingrad State Publishing House, Lenin-

grad, Russia.  (In Russian).

YURGENSON, P. B. 1979.  Biological founda-

tions of game management in forests.

Lesnaya Promyshlennost, Moscow,

Russia.  (in Russian).

Z A B L O T S K A Y A ,  L .  V . , a n d M .  M .

ZABLOTSKAYA.  2002.  Anthropogenic

effects on moose populations in the

southern taiga.  Alces Supplement 2:131-

135.

ZLOTIN, R. I., and K. S. KHODASHOVA.  1974.

The role of animals in biological turno-

ver of forest-steppe ecosystems. Nauka,

Moscow, Russia.  (in Russian).