It‘d-"‘6 ”a!

 

This is to certify that the

thesis entitled

ECOLOGY AND MANAGEMENT OF MARAL (Cervus elaphus maral)
IN NORTHEASTERN IRAN, i976'l978

 

presented by

Bahram Hasanzadeh-Kiabi

has been accepted towards fulfillment
of the requirements for

Ph.D. degree in Fisheries and
Wildlife

 

Gang, K 64:44,
J

Major professor

Date October 20, 1978

 

0-7639

. g‘ ’35:. n.“ .r: .3.

 

 

 

 

 

ECOLOGY AND MANAGEMENT OF MARAL (Cervus elaphus maral)

 

IN NORTHEASTERN IRAN, 1976-1978

By

Bahram Hasanzadeh-Kiabi

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Fisheries and Wildlife

I978

ABSTRACT

ECOLOGY AND MANAGEMENT OF MARAL (Cervus elaphus maral)
IN NORTHEASTERN IRAN, T§75-i§78

 

By

Bahram Hasanzadeh-Kiabi

Habitat and population interactions of the Persian red deer

or maral (Cervus elaphus maral) were studied in the Mohammad Reza

 

Shah National Park, Iran, during l976-1978 in the three vegetation
types present: Caspian deciduous forest, transition zone, and steppe.

Hahn's census method gave a population average of l897 over
an 18-month period for the entire park. Based on pellet group
counts there was a population of about 2096 maral in the park.

The average maral group size was 4.6. The sex ratio among
adults was 27 stags per 100 hinds. The number of calves produced per
lOO hinds was 28. The annual mortality rate was l3.9% and life
expectancy was 6.7 years.

The maral is mainly a grazer, preferring grasses over woody

forages. 0f the 19 plants eaten by the maral, Dactylis glomerata

 

was the most-preferred food, the year around. Euonymus sp., and

Colutea persica were highly-preferred winter browse species.

 

Pellet groups were correlated positively with shrub abundance.
Showing a preference for forest meadow and transition zone habitats,

the maral's food consumption was estimated to be at the rate of 2.87

Bahram Hasanzadeh-Kiabi

and 3.6 kg/day in winter and spring, respectively. The carrying
capacity of maral range was estimated to be 15.6 ha/maral and ll.0
ha/maral for transition zone and forest meadows, respectively.

Maral populations showed signs of over-abundance. There
was an excess of old animals and, in comparison with American and
European findings, the rate of recruitment was low. Of all the
maral classified, ll.0 percent were in poor condition. Wild pigs
were important competitors for food.

Leopards and wolves were the major predators capable of
preying on maral. Together, it was judged that they killed about
153 or 7.7 percent maral annually. Poaching, however, may have
caused more adult mortality than predators.

Recommendations are made for maintaining maral populations
in the national park. It is believed that the excess animals,

probably more than l60, should be removed by controlled shooting.

Tb:

Dr. George A. Petrides

ii

ACKNOWLEDGMENTS

I wish to acknowledge the cooperation of the Iran Ministry
of Science and Higher Education, Iran Department of Environment, and
Michigan State University. I am especially grateful for the kind
help of Ing. Termeh in Evin Institute of Iran. The visit of Dr. G.
Petrides during my data collection in Iran is deeply appreciated.

My principle debt is to the members of my doctoral committee,
Drs. George Petrides, Rollin Baker, Leslie Gysel, and Duane Ullrey
for their keen interest, numerous helpful suggestions, and construc-
tive reviews of the dissertation manuscript. I especially thank
Dr. George Petrides, my major professor and committee chairman, for

his guidance and encouragement.

TABLE OF CONTENTS

LIST OF TABLES .
LIST OF FIGURES

INTRODUCTION

The Present Investigation .
The Study Area

METHODS

Maral Abundance and Herd Composition
Vegetation Types . . . . .
Seasonal Use of Habitat

Seasonal Food Availability

Food Use and Preference Values

Competition for Food Between Maral and Other Large

Herbivores .
Range Condition and Trend

RESULTS

Herd Composition .
Reproduction and Mortality Rates
The Habitat and Its Use
Seasonal Use of Habitat
Seasonal Food Availability
Food Use and Preference Values
Competition for Food .
Range Condition and Trend.
Carrying Capacity .

Physical Condition of Maral
Mortality Causes .

DISCUSSION AND RECOMMENDATION .
SUMMARY

LITERATURE CITED

iv

Page

viii

NN

Table

10.

11.

12.

LIST OF TABLES

Size categories of maral hinds, Mohammad Reza Shah
National Park, Iran, 1976-1978 .

Size categories of maral stags, Mohammad Reza Shah
National Park, Iran, 1976—1978 .

Maral densities as determined by the Hahn transect
method, Mohammad Reza Shah National Park, 1976-1978 .

Average maral population densities as determined by the
Hahn transect method, Mohammad Reza Shah National
Park, Iran, 1976-1978 . . . . .

Analysis of variance in deer observation data, Mohammad
Reza Shah National Park, Iran, 1976-1978

Maral densities as determined from fecal pellet-group
counts, Mohammad Reza Shah National Park, Iran,
autumn season 1976, 1977.. . . . .

Maral population densities as determined from fecal
pellet-group counts, Mohammad Reza Shah National Park,
Iran, Autumn 1976, 1977

Group size of maral in the Mohammad Reza Shah National
Park, Iran, 1976-1978 . . . . . . .

Proportions of size categories from eight counts of
maral between November 1976 and April 1978 in the
Mohammad Reza Shah National Park, Iran .

A comparison of maral pooulations in 1975 and 1977,
Mohammad Reza Shah National Park, September 1975 and
1977.. . . . . .

Species composition of the Caspian deciduous forest
relative to Quercus castanaefolia, Mohammad Reza Shah
National Park, Iran, 1976-i978. . .

Species composition of the transition zone relative to

 

Quercus macranthera, Mohammad Reza Shah National Park,
Iran, 1976-1978 . .

Page

20

21

23

24

25

26

27

31

33

34

Table Page

13. Species composition of steppe zone relative to Festuca
ovina, Mohammad Reza Shah National Park, Iran, 1976-
1978 . . . . . . . . . . . . . . . . . . 35

14. Comparative seasonal utilization of habitat types by
maral as indicated by pellet group abundance,
Mohammad Reza Shah National Park, Iran, 1976-1978 . . . 37

15. Observations of maral by vegetation type, Mohammad
Reza Shah National Park, Iran, 1976-1978 . . . . . . 38

16. Relations between maral pellet groups and numbers of
trees, shrubs, and herbs, Mohammad Reza Shah National
Park, 1976-1978 . . . . . . . . . . . . . . 39

17. Percentage distribution of stags and hinds on wintering
areas in winter 1977 and 1978, Mohammad Reza Shah
National Park, Iran . . . . . . . . . . . . . 41

18. Seasonal changes in the average available herbaceous
forage supply in the Mohammad Reza Shah National Park,
Iran, 1976-1978 . . . . . . . 43

19. Seasonal abundances of the browse in the Mohammad Reza
Shah National Park, Iran, 1976-1978 . . . . . . . 44

20. Browse supply (kg/ha) by species in the Mohammad Reza
Shah National Park, October 1977 . . . . . . 45

21. Maral food preference ratings as determined by bite
counts, Mohammad Reza Shah National Park, Winter 1977 . 47

22. Browse consumption by 31 maral, 16 wild pigs and 2
wild sheep during a 114-day use at Karkoly Spring
feeding ground, Mohammad Reza Shah National Park,
June 1977 . . . . . . . . . . . . . . . . 48

23. Dry-weight herbaceous forages removed by 31 maral, 16
wild pigs, and 2 wild sheep during a 114- day use at
Karkoly Spring feeding site, Mohammad Reza Shah National
Park, June 1977 . . . . 49

24. Maral food preference ratings as determined by

observation, Mohammad Reza Shah National Park,
Spring 1977 . . . . . . . . . . . . . . . 50

vi

Table Page

25. Maral food preference ratings as determined by observa-
tion of wild maral, Mohammad Reza Shah National Park,
Summer 1977 . . . . . . . . . . . . . . . . 51

26. Maral food preference ratings as determined by
observation, Mohammad Reza Shah National Park,
Fall 1977 . . . . . . . . . . . . . . . . 52

27. Dry-weight forages eaten by 16 maral (6 female yearlings,
2 spike stags, 3 young hinds, 4 mature hinds, and l
fully-mature stag) during a 33-day period at Solaiman
Koshté wintering ground, Mohammad Reza Shah National
Park, January and February 1977 . . . . . . . . . 54

28. Food consumption rate by 31 maral during a 114- day spring
use at Karkoly Spring, Mohammad Reza Shah National Park,
Iran, June 1977 . . . . . . . . . . . . 55

29. Estimated numbers of major prey and predator species in
the Mohammad Reza Shah National Park, Iran, 1976-1978 . . 56

30. Proportions of the observed diet based on the number of
bites per minute for lager herbivores comprised of 8
forage species, Mohammad Reza Shah National Park, Iran,
1976-1978 . . . . . . . . . . . . . . . . 57

31. Competition for food between maral and 5 other large
herbivores, Mohammad Reza Shah National Park, Iran,
1976-1978 . . . . . . . . . . . . . . . . 58

32. Occurrence of degrees of hedging on four major browse
species, Mohammad Reza Shah National Park, Iran,
1976-1978 . . . . . . . . . . . . . . . . 59

33. Fraying behavior of maral, Mohammad Reza Shah National
Park, Iran, 1976-1978 . . . . . . . . . . . . 61

34. Physical condition of maral in the Mohammad Reza Shah
National Park, Iran, September 1977 . . . . . . . . 63

35. Prey preferences of leopards and wolves in the Mohammad

Reza Shah National Park, Iran, 1976-1978, based on a
small number of observed kills . . . . 65

vii

Figure

LIST OF FIGURES

Major vegetation types in the Mohammad Reza Shah
National Park, Iran . .

Species area curve for the Caspian deciduous forest,
Mohammad Reza Shah National Park, June-September
1977 . . . . . . . .

Species area curve for the transition zone,
Mohammad Reza Shah National Park, Iran, June-
September 1977 . . . . .

Species area curve for the steppe, Mohammad Reza
Shah National Park, Iran, June-September 1977

Regression of standard deviations on the average
monthly maral count, Mohammad Reza Shah National
Park, Iran, 1976-1978 . . . . .

Relationship between numbers per group of red deer
and their cumulative percentage frequencies on a
normal probability scale. Frequencies beyond 99.2
percent are omitted . . . . . . . .

viii

Page

11

12

13

22

29

INTRODUCTION

The European red deer as well as the maral, other Asiatic
forms and the North American elk all have been grouped in the species

Cervus elaphus. The range of the maral (Q, g, maral) includes the

 

Caspian provinces of northern Iran, Crimea, Asia Minor, and
Caucasus.

g, elaphus is a characteristic inhabitant of the deciduous
forest regions of Eurasia and North America. It is also found in
the Mediterranean chapparal. Mature stags in eastern Europe have
been recorded with antlers over 130 cm long and weighing more than
12 kg. The largest pair of red deer antlers known weighed 19 kg
(Corbet and Southern, 1977). In early spring, the stag's antlers
are shed and six weeks later new antlers begin to grow. By the end
of May, they are full size (Whitehead, 1950).

Red deer are herd animals, with the sexes separate most of
the year. Female herds are made up of hinds, calves and include
stags up to two years old. The stag herds consist of older males
(Harris and Duff, 1970). With the mating season, or rut, the stag
herds break up, usually in October. Each mature stag attempts to
move into an area occupied by a herd of hinds. Under normal wild
conditions, estrus probably first occurs in the third year of a hind's
life (Chapman, 1974). Following a fertile mating, gestation occupies
about 250 days (Chapman, 1974). The female does not produce a calf

1

every year (de Nahlik, 1974), especially in poor environments
(Harris and Duff, 1970). In early summer the calves are born,
usually singly. The weight of a newborn calf can vary between 5 and
10 kg (Harris and Duff, 1970). Full adult size is not attained
before the 7th year (Lowe, 1971). Maximum longevity in captivity

is about 20 years (Corbet and Southern, 1977).

Home ranges are variable in shape but in Scotland, on the
island of Rhum, they involve areas of about 400 ha for a hind and
800 ha for a stag (Lowe, 1969). There is reason to believe that the
farming of red deer for meat production will prove to be technically

feasible (Blaxter et a1., 1976).

The Present Investigation

 

An 18-month study of the maral or Persian red deer was
started at Mohammad Reza Shah National Park in November 1976. Some
findings from a 4-month investigation during the summer of 1975 also
contributed to the effort.

Objectives were: (1) to determine the population character-
istics and environmental interactions of the maral; (2) to assess the
carrying capacity of the range for this large deer; (3) to describe
interactions between other animals and the maral; and (4) to determine
how maral abundance affected wildlife and wilderness standards in the

national park.

The Study Area

 

The Mohammad Reza Shah National Park was the first area in

Iran to be designated as a national park. Located in the province

of Khorasan from 37° 36' to 37° 16' north latitude and 56° 17' to
55° 44' east longitude (Figure l), the terrain is mountainous with
the altitude varying between 380 and 2819 meters. The park is about
126,000 hectares in area (de Vos et a1., 1977).

2.an ._m .9 wo> mu ES“: 69. .de .95sz
£25 «Nwm 38620.2 9: E 89: cozflmm? 8.3.2 .P 2:9”.

”#2300 nSOtQ OT—
km44m11m<mt< 53.5% 0
NEW uz‘tmhzg.
MFImOXIZ<E‘<JOm a "I
whommZQQH wi— ExO. n 0

295.5191 nm<mm< Sim
mammpm 30.. H

15.6 “.540

         

    

   

mammem :9: D ,2 .593 \\ \
Samoa 23.55 S i 4:3”...umV. ”with? \
mzoN 20:3qu mm 23% . A!«we:mrfiwmimfi
.. film/«Win... .2.me .X¢J.m.v1
. /rUV..MV/¢/ . fir...
. . 2;. as? a
.... .. . av OHNIWO— .L .
_
3:35. :98 ,.
o o
\ omuousoom
ouumz<z<xo

Z<Pw<x

METHODS

Maral Abundance and Herd Composition'

 

The Hahn (1949) deer cruise-line technique was one of two
methods used to enumerate the maral herd. Transects were laid out
randomly in each of the major vegetation types (Figure l). The
length of the transects varied because of topography. Prior to the
census, distances to which maral could be seen were measured by
pacing at right angles on each side of the center line at loo-meter
intervals along the transect. The average of these visibility
distances multiplied by the length of the transect gave the area
censused.

A survey was conducted each fall and winter. These were
accomplished on November 27 through December 12, 1976; March 1 to 12,
1977; November 1 to 12, 1977; and February 7 to 19, 1978. Two counts
also were made each spring and summer following the same routes.
These were completed April 4 through May 3, 1977; May 30 through
June 9, 1977; August 27 through September 8, 1977; and September 13
to 24, 1977. Maral observed on the transects were tallied according
to size and age criteria (Tables 1, 2). Specimens for which sex and
age could not be determined with certainty were placed in an
unclassified category. In each of the two census methods, popula-
tion densities were computed by averaging the density estimates on

all transects used.

TABLE l.--Size categories of maral hinds, Mohammad Reza Shah
National Park, Iran, 1976-1978.

 

Size Categories

Descriptions*

 

Calf

Very young hinds
(yearlings)

Young but fertile
hinds

Fully-matured
hinds

01d hinds

Spotted

Smaller in the shoulder than the older animals,
in a herd yearlings normally trail behind the
most recent calf which follows the mother.

Up to the age of 2 years, the short length

of the head, which assumes full size by the
age of 2% years, is a good indicator of age.

Aged 3-5 years, head and body start to fill
in.

Body looks sturdy and well-filled, ears
appear firm and fully responsive to animal
reflexes. Animals normally will not show
their ribs other than perhaps at the end of
winter. Aged 6-10 years.

The neck gets thinner. The ribs are visible
through the skin for most of the year. The
entire head appears longer and more emaciated.
The ears become flabby. Aged over 10.

 

*From de Nahlik, 1974.

TABLE 2.--Size categories of maral stags, Mohammad Reza Shah
National Park, Iran, 1976-1978.

 

 

Size Categories Descriptions*
Calf Spotted
Spike stag- Small head, short bo y, long egrs,simp1ep
yearling antlers without brow and trey tines.

to 2 years.
Young but Antlers with bey524and brow1 tines. Trey3
fertile stags are about to fork. 2- 5 years old.
Fully mature The body looks sturdy and well-filled,
stags antlers develop a crown.5 6-10 years of age.
01d stags Antlers deteriorate, tines are shortened,

crown disappears. Over 10 years.

 

*From de Nahlik, 1974.

l

Brow

2Bey =

3

4

5

Trey

Fork

Crown =

The lower point on the antler-~sometimes called the
first antler.

The second point or tine of a stag's antlers.
The thired point or tine of the antlers.

The two points on the extremity of the antlers in
the form of a fork.

The uppermost cluster of three points of a red deer
head, if they form a crown.

Transects were distributed so that all parts of the densely
populated range was represented. They were spaced further than 3 km
apart, however, to prevent counting a maral more than once. A11
census routes followed existing trails in order to reduce noise
caused by the observer. From a statistical standpoint, locating
transects randomly throughout each area being censused would have
been desirable. Because of noise caused by the observer cruising
through the vegetation, however, that procedure would have reduced
the numbers of maral seen and seriously biased the results. Clearing
new paths would, in addition, have modified the vegetation pattern.

The assumptions had to be accepted that: (l) the sample
counts were equivalent to random samples and (2) the number of maral
observed equalled the total number of maral present along the
transects surveyed. The analysis of variance for deer counted was
calculated on the basis of the number of maral seen per 1000
hectares. Because the standard deviation of counts within a month
tended to vary in proportion to the average monthly maral count, the
variate transformation Y = log (X + 73.10) was calculated and used
in the analysis.

Using binoculars, care was taken not to disturb the deer.
Most observations were near dawn and dusk when maral were most
active. Notes were made also of other animals and their signs seen.
This was in an effort to determine possible competition and other

interactions with maral.

Pellet-groups were counted as an alternate method to deter-
mine maral density. The 4 m2 circular plot (Riney, 1957) was used.

Sample plots were distributed randomly in areas of high-
intensity use where it has been found to be more efficient (Neff,
1968). Few pellet-group sampling systems have employed a completely
random distribution (Neff, 1968). Leopold et a1. (1951) stated that
best estimates of deer numbers in the California mountains were
obtained from transects which were distributed by subjective selec-
tion and which followed trails or ridge tops.

In the present study, the number of plots needed at each
sample site was determined from preliminary surveys applying the

formula used by Grieb (1958):

N = (t 0.10)2 s2
(0.20 x T)2

where:
N = number of plots needed
52 = variance in pellet density from preliminary data
X = mean pellet density based on preliminary data

0.20 = selected risk of error expressed as a fraction of mean
pellet density

t 0.10 = tabular value of t for a selected level of probability.

Based on more or less randomized preliminary pellet-group
surveys, 113, 284, 379, and 115 sample plots were calculated to be

necessary in steppe, transition zone, Caspian deciduous forest, and

lO

forest meadows, respectively. The distance between plots was
decided on an arbitrary basis to be 10 meters. Pellet-group counts
were conducted only in the autumns (of both 1976 and 1977) since the
lush vegetation growth in spring and summer tended to hide pellet-
groups at those seasons. Pellet-group counts were also conducted

in late winter and early spring on a few wintering grounds while

gathering data on maral food consumption and preferences.

Vegetation Types

 

Three major vegetation types were identified as the Caspian
deciduous forest, a transition zone, and the steppe. Species compo-
sition was determined on sample-plots whose sizes were determined by
the nested plot technique (Mueller-Dombois and Ellenberg, 1974).

For the three vegetation types, the areas were 200 m2, 40 m2, and 4 m2
for the Caspian deciduous forest, transition zone, and steppe,
respectively (Figures 2, 3, 4). Choosing accuracy and t values which

seemed appropriate, the number of plots needed for each vegetation

type were determined by the de Vos and Mosby (1969) formula:

n = szztz .05
d .10
where:
n = number of plots needed
52 = variance in preliminary data
d2 = (0.10 x 'x')2

7'= mean of the prelimonary data

11

.280ch m 2 .33 m_
3385 86on 9655 $5 .03 .mEEE 2 8.6983

11
O

m o. 3:93 Emucm. o

.3255: 3.0ch c. 338:. can a 22>
2:03 «So c. 388:. $3 m 29.3 32m :8an m

83?. no.6 2.93:6 .0 22a 5 952 “203» .o 235:: u < 6:33

.28:
.mnEoEow - 92:. .{ma .98sz :95 Sum omEEmcoz
.592 253208 .5330 9: .2 misc new - 3625 .N 930E
.mmwhuz wm<aom . mmflm ._.o.E
com. com 00¢ Dow 00.
_ _

 

N

'0

V‘

 

 

.LO'ld 83:! 33103135 :10 838101le

12

83 P .mnEoEmm - 0:3. .cm: £ch .95sz :95
3mm 368205. .chN .5293: 05 .2 misc coca 38on .m Saul

Ammwsz mm<30m v mmflm ._.o._n_
0mm 0mm O¢N CON Ow. ON. om O¢

 

 

 

q_fl________‘__d ‘
_
_ «N
_ n
_ now.
_ em.
_ a
_ mm
_ kw
_ mg
_ 19a
_l___.nlw.
IN.
9 * Im—

 

13

.29 .ooEmEmw - 0:3. 68. .{md .95sz
:95 «Non. 3552.22 .2506 9: .2 02:0 «Em - 8.025

.mmmhmz umdfiom. mmN.m Four...
6. v. N. O. m w

.v 9:9“.

 

V
a _ _ _ _ _ a
_

 

MN.
__

_

 

.LO‘ld 83d SBIOBdS :10 338111011

14

0.10
t 0.05

designated accuracy

tabular value of "t" for the selected level of
probability.

Based upon the formula by de Vos and Mosby (1969), it was
determined that sample sizes of 126, 114, and 118 plots were
necessary for the Caspian deciduous forest, transition zone, and
steppe, respectively. Within these plots, the numbers of living
forbs, grasses, shrubs, and trees were tallied.

Frequency, density, and index of species associated were
determined for each of the three major vegetation types. Absolute
frequency refers to the number of plots in which a given species
occurs. For comparing different plant communities, it is convenient
to convert absolute frequency into relative or percentage frequency.
Density refers to the total number of individuals per plot. The
index of species association (IAp) is used to evaluate the correla-
tion in presence and absence of species in a vegetation type, the
floristic composition of the type, the distribution of the species
therein, and their ecological relationships (Mueller-Dombois and

Ellenberg, 1974).

Seasonal Use of Habitat
An index of habitat use by maral was calculated from pellet-
group deposition over the three major habitat types plus forest
meadows. Moran's utilization index (1973) was used to compare pellet

group deposition between habitat types.

15

Percentage of_pellet groups in type

Ut111zat1on 1ndex = Percentage of area occupied by type

Habitat use was measured in winter and in summer. Winter was
identified as the 100-day period following the date of heaviest
leaf-fall (November 19, 1976 and November 16, 1977) and extending
through snow-free days in spring. Summer use was arbitrarily chosen

as an indeterminate period prior to leaf fall (September, 1977).

Seasonal Food Availability

All plants within clip-plots, each 4 m2 in area and located
randomly in maral habitat, were sheared to a height of 1.8 meters
to assess dry-weight forage composition and production. Specimens
of both woody and herbaceous forages were randomly selected from
different plants to determine mean dry weights for the uneaten plants
of each species. All plants were dried in a standard laboratory
oven at 105°C for 24 hours before weighing.

Measurements of acorn crops were made in various sections
of the park because of the importance of acorns as a wildlife food.
Measurements of acorn crops were made by counting acorns on sample
branches, and by determining total number of branches per tree for
the two different species of oaks. Counts of the fruits present
were made before the acorns dropped, using binoculars. Mean dry
weights for acorns were determined after treatment in a standard

laboratory oven at 105°C for 24 hours.

16

Food Use and Preference Values

 

Shafer's (1963) twig count method and the dry-weight differ-
ence method of Beruldsen and Morgan (1934) were employed to determine
the availability and ungulate use of woody browse and herbaceous
forage.

Food habit studies of the maral were carried out at Karkoly-
Spring (Figure 1). The sharp cuts made on vegetation by hares and
mice were distinguishable from the broken vegetation induced by
feeding ungulates, which lack upper front teeth. It was thought,
therefore, that signs of the field mouse (Apodemus sylvaticus),

social vole (Microtus socialis), Afghan mole vole (Ellobius

 

fuscocapillus), and long-tailed hamster (Calomyscus bailwardi) were

 

 

not misidentified as deer food removals. Winter food habits studies
of the maral were carried out at Solaiman Koshte (Figure 1) where an
area of approximately 380 ha supported a population of 16 maral.
Only maral used this site. Winter food habits were determined by
locating a fresh maral track and recording the signs of fresh
browsing along its trail. Each browsed twig was designated as one
instance of use. The number of twigs taken and the number of twigs
available within one meter of each side of the trail were recorded
by species. Besides the exact counting of bites on vegetation, a
description of the habitat, snow depth, crust condition, and length
of trail were recorded. To distinguish recent bites from old ones,
fresh twigs were broken frequently and the color and condition of

the broken surfaces compared. Shortly after a fall of snow, fresh

17

trails were followed backwards to avoid disturbing the animals. Each
bitten twig was recorded and where tree bark was gnawed, the height,
thickness, and area consumed were measured.

Though the percentage of each species utilized also indi-

percentage in diet

cates comparat1ve forage preferences, the rat1o percentage available

yields a figure which, if over 1.00, indicates relative degrees of
preference and if under 1.00, reveals degrees of unattractiveness
(Petrides, 1975).
Competition for Food Between Maral
andFOther Large Herbivores
Feeding records were obtained for the wild pig (Sus scrofa),

goitered gazelle (Gazella subgutturosa), wild sheep (Ovis ammon),

 

wild goat (Capra hircus aegagrus), and roe deer (Capreolus capreolus)

 

as opportunity afforded. Diets of these larger herbivores were
compared for similarity with that of the maral, being expressed as
the degree of overlap. "Overlap" was used in the same sense as Horn
(1966) and calculated by summing the fraction of each herbivore's
recorded diet which had counterparts in the diet with which it was
being compared. The extent of habitat overlap also was estimated
based on range use and preference as evidenced by field observation.
The term "competition coefficient" is proposed and quantified as the

product of diet-overlap and range-overlap fractions.

Range Condition and Trend
The effects of maral on vegetation were observed along the

routes followed while making pellet-group counts. The three shrubs

18

nearest to each 4 m2 plot were examined and the proportions of these
plants which were hedged in varying degrees were recorded. The
shrubby species available to maral were classified into five groups

in the manner of Riney et al. (1959):

0 = No evident browsing effects

1 = Shrub lightly hedged, shape only slightly altered

2 = Plant hedged, but recovering or capable of recovering
from browse pressure

3 = Shrub size held entirely in check by animals

4 = Plant killed by maral.

RESULTS

Population Densities

 

Maral densities per 1000 hectares were calculated by the
Hahn method for each vegetation type and season (Table 3). Combining
these densities with the areas of each cover type yielded an overall
number of 1897 maral present on average during the lB-month study
period in the entire park (Table 4).

When variance in the number of maral seen per 1000 hectares
of land was appraised, there was an evident tendency (Figure 5) for
the standard deviation of counts within a season to vary in propor-
tion to the average seasonal maral count. Transforming the data
(Table 3) by the variate Y = log (X + 73.10),* differences in deer
densities between seasons, areas, and their interactions (Table 5)
were indicated to be highly significant.

Using the fecal pellet-group data with a defecation rate of
10 pellet-groups per day for red deer (Riney, 1957), the mean autumn
population estimate was 2096 deer (Tables 6 and 7).

Herd Composition

 

Average maral group size in the park was 4.6 f 1.1 (Table 8).
Female deer outnumbered males by more than 3 to l. The average sex

ratio observed among adults was 27 stags:100 hinds (Table 9).

 

* a
Based on Hartley et a1. (1955); Y = log (X + E).

19

20

 

 

 

 

 

mm um um .m we mm mm mm wu<mm><
mm mm .4 mm. mm mm A. a. so..a> »_oxam¥
Lona: use omcpeuwcwmcmemgu
"UZON onhHmz<mh
om mm m. we 4. mm as am oomgm< - .gmmaaa>am
mm m AN AN .0 am mm m moaox - .mpgxae
mo. we m so mm 48 on. m cmcmaNom
an no .__ «A NN. mm aw .m cmgmooucagx
am me an o“ em w. m“ mm .Pao._< - mm_aae<
“mzoo<uz emmzoa
mm ._ o. m. a. A mm o ._o;mauo;¥ -.moa.
am e o. o. m m. .4 N. mmpgmwsacu - cama=u<
”Amaze. m=o=a.umo z<.am<o
mum. Rum. “Am. Aka. Rum. hum. Num— who.
empcwz ppm. seesaw gossam mcwgam m:_egm empcwz ppm.

 

mmgmpum: coop Lea ammo we gmasaz

ao.m

 

chowuwz cmgm mNmm umesmsoz

.wnmpumm¢p .xgma

.vosums «ommcoga ccm: mgu an cmcwEngmu we mowuwmcmu Pmemz-u.m m.m<h

21

TABLE 4.--Average maral population densities as determined by the
Hahn transect method, Mohammad Reza Shah National Park,
Iran, 1976-1978.

 

Average Maral

 

Vegetati°n Type SEEialfls Igggsgggtgfigs v232¥§ii§3”r§3§
Caspian Deciduous 34,100 19 648
Forest
Forest Meadows 3,400 55 187
Transition Zone 14,000 68 952
Steppe-Border* 4,500 ** 110**
Open Steppe 70,000 0 0
TOTAL 1 26 , 000 15 1TB;-

 

*
Area adjacent to transition zone (1.5 km wide)

**Not surveyed by the Hahn method. Observations of maral
over the 15-month period, however, showed that 5.8 percent of all
deer seen were in the steppe-border area (see Table 15).

22

. £372.?
68. .mea .2282 :95 «Now. umEEmzos. .ano .EmE

 

:55... 3295 9: :0 20.3.5“. Emucmfi .0 3.3931 .m 2:9“.
m .mo<mm>< 5:50:
we ow No 3‘ mm mm Cu m. N. w v
. . _ _ . _ . _ . . _
1 ¢
1 m
1N.

 

 

S ‘SlelOO A'IHlNOW JO NOIJMABCI OBVGNVLS

TABLE 5.--Analysis of variance in deer observation data, Mohammad
Reza Shah National Park, Iran, l976-l978.*

 

 

df 55 ms F
Season 3 .08 .0297 5.68**
Areas 7 .44 .0628 l3.36**
Season x Areas 21 .25 .0119 2.53**
Error 32 .15 .0047
mm 2; I;

 

*Transformed, where y = log (X + 73.10)

**Significant at the 1 percent level.

24

TABLE 6.--Maral densities as determined from fecal pellet-group
counts. Mohammad Reza Shah National Park, Iran,
autumn seasons 1976, 1977.

 

Deer per 100 hectares

 

 

 

 

 

 

 

Site
Fall 1976 Fall 1977

CASPIAN DECIDUOUS FOREST:
Adnasad - Ghamishlee ll 23
Late - Khodagholi ll 10
Gharangine - Jangle 32 22
ngEST MEADOWS:
Afralee - Alidali 14 50
Khandooshan 26 25
Bozaghan 24 4o
Takhte - Korda 63 25
Sararbaghi - Aghsoo 42 29
TRANSITION ZONE:
Gharangine - ridge and upper

Karkoly Valley 73 79
STEPPE - TRANSITION ZONE ECOTONE:
Takhte, - Almeh 39 52
OPEN STEPPE: 0 0
Average* 33.5 35.5

 

*Open steppe is not included

(No. of pellet-groups/lOOO ha)

(No. of days since heaviest

leaf fall x 10 groups/day/
maral)

Heaviest leaf fall were recorded on November 19, 1976 and
November 16, 1977.

Number of maral per 1000 hectares =

25

TABLE 7.--Mara1 population densities as determined from fecal pellet-
group counts, Mohammad Reza Shah National Park, Iran,

Autumn 1976, 1977.

 

. Average Maral
Area 1" Density per

VGQEtatTOn Type Hectares 1000 Hectares

Population for
Vegetation Type

 

Caspian Deciduous 34,100 18
Forest

Forest Meadows 3,400 34

Transition Zone 14,000 76

Steppe-Border 4,500 45

Open Steppe 70,000 0

TOTAL 126,000 16.6

714

116

1,064

202

 

2,096

 

26

TABLE 8.--Group size of maral in the Mohammad Reza Shah National
Park, Iran, 1976-1978.

 

 

2:22P 4:35:22, mazes;
1 124 20.4 20.4
2 126 20.7 41.1
3 86 14.1 55.2
4 66 10.9 66.1
5 32 5.3 71.4
6 51 8.4 79.8
7 27 4.4 84.2
8 22 3.6 87.8
9 15 2.5 90.3
10 15 2.5 92.8
11 14 2.3 95.1
12 9 1.5 96.6
13 3 0.5 97.1
14 4 0.7 97.8
15 4 0.7 98.5
16 2 0.3 98.8
17 2 0.3 99.1
18 1 0.1 99.2
19 1 0.1 99.3
23 1 0.1 99.4
49 2 0.3 99.7
58 1 0.1 99.8
72 1 0.1 99.9

 

. 1 11’ d
Average group s1ze = 5135913335 E211133= 4.6

27

 

 

 

 

mo. mo. mm. mm. mp. mc. us. we. co. mmwm mmnmnmp m.<k¢h
0 Fe. m—. Re. om. Fo. mo. Po. .0. en. wmmp gmucmz
mo. o mm. mm. m_. Fo. wo. co. co. mom nmm— P~mm
«p. mo. mp. mm. mp. co. no. mo. «0. com “nap Lwnsmunmm
cp. co. mp. mm. m—. we. no. mo. #0. nmm Rump Losszm
pm. 0 PF. mm. Fm. po. mo. mo. mo. com Rump acmgam
o mo. mm. mm. up. co. no. mo. mo. upop mnm— Lousy:
Po. o NN. mp. mm. o No. mp. OF. mmp onmp ppm;
Np. op. mm. op. mm. No. No. mo. mo. amp mnmp LmnEmuawm
mvcw: mmmum

 

.cmt. .xaaa .m:o.omz swam mNQm umEEmgoz ago a. mum. ..La<
was ono. amnew>oz cmmzpmn Poems mo muczoo ucmwm seem mwmgommumo mNPm mo mcowueoaoganu.m m.m<h

28

If the cumulative herd-size frequencies were transferred to
a normal probability scale and plotted against numbers per group then,
the frequency distributions, if normal, would form a straight line.
Actually, the data form approximate straight lines (Figure 6)
between points 3-12 and 12-18.

The data from chamois (Rypicapra rupicapra) and red deer

 

(Cervus elaphus) were bimodal (Caughley, 1964). The present data

 

and those populations indicate that group size does not affect the

animal's urge to join it.

Reproduction and Mortality Rates

 

The first calls of rutting male maral started as early as
September 2 and continued through October 5. The number of females
gathered by a breeding stag varied from 2 to 19. There was an
average of 10 hinds per harem in the transition vegetation zone and
5 hinds per breeding stag in the Caspian deciduous forest.

Gestation in red deer has been reported to last about 8
months (de Nahlik, 1974). In the park, it was confirmed that gesta-
tion approximately lasted from about September 2, when the rutting
season started, to May 10 when the calving season began, a total of
250 days.

Single calves were the rule. Only two instances of twins
were seen. In 1977 the first report of a newborn calf was May 10.
From the monthly distribution of 16 births between the years 1840
and 1870 published by Zuckerman (1953), the mean date of birth and
standard error were determined as explained by Caughley (1977) to

be 30 June f 2 days.

29

.0 0.03 000. 003.50
0.0 E0000: «.3 0:0»00 00.050000“. 0.000 5:30:03
.250: 0 :0 00.05000: 000E020: 050.3an :05
0:0 .000 00.. .0 0:03 :0: 200:5: 5020: 2:05:20: .0 052“.

m.mmmmmmmmomomowovONo.nN_

 

_______d4_du_.._

<3 01 o h to ID 0 IO N -
ant-IO 83:1 1.438711le

FOODQ’TON-

 

30

The number of calves produced per 100 hinds based on summer
data was 28 (Table 9).

Despite limited available data from the summer of l975, a
comparison between p0pulation numbers of l975 and l977 was made
(Table 10). Census results for September 20 through September 30
were available for comparison and the Mann-Whitney U-test (Elliott,
l977) was computed. The U values of 5 and 10 suggest that the two
populations were not different in density at the 5% level of sig-
nificance. In order to gain at least an approximate measure of some
maral population characteristics, it may be assumed that the popula-
tion is stationary. If so, therefore, population mortality rate and
life expectancy can be derived from the ratio of juveniles to adults.
Where j is the number of O-l year old animals in a sample of n
individuals drawn from a stationary population immediately after a
restricted annual season of births, the population annual mortality
rate is equal to the proportion of juveniles (j/n) and life
expectancy in years is (2n-j)/2j (Caughley, 1967). Based on summer
data (Table 9), the park maral had a yearly mortality rate of l3.9%

and a life expectancy of 6.7 years.

31

TABLE lO.--A comparison of maral populations in l975 and l977,
Mohammad Reza Shah National Park, September l975 and

 

 

 

 

 

l977.
September September
l975 l977

Proportions of Size Categories:

Calves .l2 .l4

Hinds .71 .66

Stags .l7 .20
Maral Seen per Census: l7, 8, 68 81,72,2,22,18
Mann-Whitney U-test:*

Rl, 2 ll 25

Ul, 2 5 10

Nl, 2 3 4

 

*

Where the calculated levels of U exceed the tabulated
values of u, the mean levels of the two samples are accepted to
be identical. U is not significant, therefore, at the 5 percent
level of significance.

32

The Habitat and Its Use
Vegetation Types
In the 37,500 ha Caspian deciduous forest dominant trees

were: Quercus castanaefolia, Parrotia persica, Carpinus betulus,

 

 

 

and Crataegus s2, (Table ll). Man-caused forest meadows covered
8.7% of this area and were composed of (l) grasses 59.9 percent by

stem count, including Avena sativa, Dactylis glomerata, Bromus

 

 

briziformis, Bromus sterilis, Bromus japonicus, Bromus gracillinus,

 

Hordeum bulbosum, and Aegilops tauschii; (2) legumes l3.8 percent

 

 

in which Vicia persica, Vicia sativa, Medicago orgicularis,

 

Medicago polymorpha, and Trifolium angustifolium were prominent;

 

 

(3) other forbs l5.9 percent, mainly Achillea nobilis, Tragapogon

 

graminifolius, Phleum paniculatum, Stachys olympica, Carex

 

sylvaticum, Scabiosa micrantha; (4) shrubs 9.7 percent Rubus sp.

 

and Rosa beggeriana; and (5) ferns 0.7 percent, mainly Pteridium

 

aquilinum.

The transition zone, of approximately l4,000 ha, was
characterized by shrubs and scrub trees, common among them were
Acer ibericum, Paliurus spina-christi, Colutea persica, Quercus
castanaefolia, Quercus macranthera, and Berberis vulgaris (Table l2).

dominated by Festuca ovina and Artemisia sp. (Table 13). Broad-

 

leaved forbs were frequent but trees (Acer ibericum) were present

mainly in ravines and in the forest border ecotone.

TABLE 11.--Species composition of the Caspian deciduous forest

33

relative to_Quercus castanaefolia, Mohammad Reza Shah
National Park, Iran, 1976-1978.

 

 

 

 

Frequency* Density**

Species IA ***

Absolute Percent Sum Mean
Quercus castanaefolia 126 100+ 573 4.5 100+
Parrotia persica 77 61 225 2.9 61
Crataegus sp. 66 52 181 2.7 52
Colutea persica 66 52 167 2.5 52
Prunus caspica 63 50 169 2.7 50
Paliurus spina-christi 57 45 137 2.4 45
Mespilus germanica 54 43 154 2.8 43
Carpinus betulus 52 41 189 3.6 41
Rubus ulmifolius 39 31 86 2.2 31
Zelkova carpinifolia 28 22 73 2.6 22
Quercus macranthera 13 10 63 4.8 10
Vitis silvestris 10 8 25 2.5 8
Acer velutinum 7 6 15 2.1
Diospyros lotus 5 15 2.5

 

*
Number of plots in which a given species occurred.

**

Numbers and percentages of individual.

**

*
Index of species association as compared Wlth Quercus.

+Due to the 100 percent constancy for Quercus, values for
its percent constancy and IAp were identical.

IA
where c =
together;
a:
occurred alone; and
b:

found alone.

P

- a + b + c
the number of plots in which the two species occurred

C

x 100

the number of plots in which one of the two species

the number of plots in which the other species was

34

TABLE 12.--Species composition of the transition zone relative to
Quercus macranthera, Mohammad Reza Shah National Park,
Iran, 1976-1978.

 

 

 

 

 

Frequency* Density**
Species IA ***
Absolute Percent Sum Mean p
Quercus macranthera 101 89 161 1.6 100
Paliurus spina-christi 97 85 358 3.7 78
Colutea persica 97 85 272 2.8 78
Quercus castanaefolia 97 85 227 2.3 73
Berberis vulgaris 81 71 149 1.8 60
Acer ibericum 86 75 166 1.9 65
Crataegus sp. 64 56 88 1.4 50
Lonicera floribunda 53 46 103 1.9 42
Cotoneaster sp. 49 43 95 1.9 41
Rhamnus sp. 35 31 68 1.9 31
Euonymus sp. 34 30 77 2.3 25

 

*
Number of plots in which a given species occurred.

**Numbers and percentages of individuals.

***
Index of species associated as compared with Quercus.

35

TABLE 13.--Species composition of steppe zone relative to Festuca
ovina, Mohammad Reza Shah National Park, Iran, 1976-1978.

 

 

 

 

Frequency* Density*

Species IA ***

Absolute Percent Sum Mean p
Festuca ovina 84 71 562 6.7 100
Artemisia absinthium 68 58 547 8.0 56
Dactylis glomerata 67 57 214 3.2 47
Agropyron cristatum 47 40 172 3.7 46
Teucrium polium 57 48 135 2.4 44
Centurea bahen 49 41 154 3.1 40
Ephedra sp. 61 52 475 7.8 39
Galium tricorne 50 42 115 2.3 36
Poa bulbosa 42 36 126 3.0 33
Colutea persica 42 36 52 1.2 31
Acer ibericum 32 27 36 1.1 25
Juniperus sp. 18 15 18 1.0 19
Prunus sp. 18 15 21 1.2 15

 

'11:

Number of plots in which a given species occurred.
1H:

Numbers and percentages of individuals

*** . . .
Index of species assoc1ation as compared with Festuca.

36

The Caspian deciduous forest, the transition zone, and the
steppe zone were judged to be ggercus-Parrotia-Carpinus, Quercus-

Paliurus-Colutea, and Festuca-Artemisia associations, respectively.

 

Seasonal Use of Habitat

Utilization index values (Table 14) indicated that maral have
both winter and summer preferences for meadows in the Caspian
deciduous forest. It is interesting that 5.8 percent of the maral
seen (Table 15), however, were in steppe vegetation. Their pellets
were found even in the Almeh Valley, 3—4 km from the woodland.

Maral appear to have been numerous on the Caucasian steppes
during the Pliocene (Verseshchagin, 1967) and to have occupied a more
widespread range then than today. During recent times, major causes
of maral reduction, according to Verseshchagin (1967), appear to have
been a combination of competition from livestock and indiscriminate
hunting. It is possible that stag hunts by horsemen forced the
retreat of maral into the forests where they were out of reach of
mounted hunters. Today in spring and summer deer tend to migrate
from forests to the open steppe in escape from blood-sucking insects
(Verseshchagin, 1967).

Shrub abundance seemed to be the most important vegetative
factor associated with high pellet group density and mixed trees and
shrubs was the vegetation type most preferred by maral (Table 16).

The number of pellet groups was considerably higher where
numerous shrubs were associated with trees than where plots contained

trees and few or no shrubs.

37

. 00NH :w mqsogm pmppmniw n

0050 an umwaauoo 00:: x u

 

A:.=v xaaew eovuanpppzt

 

 

 

o o o o m.mm o o maampm cage
0:0uoum 0:0N

om._ oo.p m.m m.m m.m mam o cowuwmcmshuwaamam

mm.e No.p «.me m.__ o.FF mm: m_ 0:0N compwmcmgh

No.5 mm.~ ¢.mp m.o ¢.N mmm mm mZoomwz pmmgou
pmmcou

oo.P mw.~ o.- N.wn o.n~ em :0 msoszumo :qummu

*H-: 3H1: umppma 00__0¢ 00:< gossam gmpcwz
cmesam gmucwz 0:00:00 0:00:0a pcmugmm 00a» powwow:
gmeszm :0»:w3 0: :0:

manage umppma 0mm:0><

 

.mmmpnmnmp .:0LH .xgma ~0:0wumz gmsm mem 0:250:02 .mucmvcznm

asogm uwp_0q x: umpmuwccw m0 pmgms an mmaxu umu_nm; mo :oquNPFPuz chommmm 0>wumgmasouu-.ep m¢m<h

38

.e 0pnmh 00m:

 

 

 

 

 

 

 

o.oo_ ovpv o.oop o.oop o.oop o.oop mmmp mum «mop 0mm 4<pop
«m.m .em m.NF o.m ¢.o n.m mup mm m mm maamum
o.m~ mmop _.¢¢ F.mm w.o m.~¢ mp: map amp mop 0:0N :0wummcmgh
w.mm Npmm ¢.mm n.00 w.Pm _.me mmm mnp amvp wNF mzoummz ummgom
umwgo:
m.- mmm o.m m.m o._m e.m ox Pm ope Fm mzozcwumo :0wammu
&
mnmp . mxmp cmsszm mcwggm gmacwz Ppmd gmsszm .mcvsam qucwz ppm:
00»» :0wpmpmm0>
punch :00m page: 00 mwmmacmugma :00m Fmgmz 00 :00532

 

.mmmpumnmp

:mcH .xgm: Pacemamz gmcm m~0m 0:550:02 .maxu :owumumm0> an Fugue 00 m:o_um>gmmaonu.m_ u4m<h

39

TABLE 16.--Re1ations between maral pellet groups and numbers of
trees, shrubs, and herbs, Mohammad Reza Shah National
Park, 1976-1978.

 

 

Mean Pellet Groups Trees* Shrubs** Herbs***
Per Plot No. per ha No. per ha kg/ha
0.6 0- 25 216 529
0.8 26- 50 362 451
3.2 51- 75 412 286
4.9 76-100 448 255
1.3 101-125 232 107
1.2 126-150 132 78

 

*
75 percent Quercus castanaefolia and 25 percent other
species.

**50 percent Quercus castanaefolia, 20 percent Colutea
persica, 20 percent Prunus caspica, and 10 percent other species.

 

*1: .
* 45 percent Bromus sp., 25 percent Agropyron trichophorum,

10 percent Medicago sp., and 20 percent other species.

40

The two consecutive winter seasons during which studies were
carried out differed considerably from each other with respect to
the depth and persistence of snow. It seems that snow depths of
about 45 cm seriously impeded the movements of maral. Two days
after a snow of more than 45 cm., marals had moved to the Solaiman
Koshte (a transition vegetation type) wintering ground at a lower
elevation.

Maral stags tended to associate with hinds mainly during the
rutting season. For most of the year they lived in segregated herds.
Although it has been known (Darling, 1937) that stag leave the hinds
after the rut and move to different home ranges, it evidently has
not been published that such spatial segregation occurs on common
winter ranges. Stags outnumbered hinds in winter at high elevations
(Table 17, Upper Karkoly Valley) and are thought to be more abundant
at other seasons as well. This type of habitat segregation has also
been recorded for wapiti (Flook, 1970) and bighorn sheep (Geist and
Petocz, 1977). In winter, the females and young stayed at lower
elevations where the weather was milder, snow-free days were more
frequent, and grasses were more abundant. As Geist and Petocz (1977)
have pointed out, it is possible that spatial segregation of the
sexes benefits hinds and calves and, therefore, maximizes herd

reproductive fitness.

41

TABLE 17.--Percentage distribution of stags and hinds on wintering
areas in winter 1977 and 1978, Mohammad Reza Shah

National Park, Iran (n =
observations).

1984, including repeat

 

 

 

 

 

 

 

Approximate -
. Stags Hinds
Census Area Altzgudes Percent Percent
Caspian Deciduous Forest:
Gharange Jangle 1500 17.07 3.97
Golestan - Golshan 500 2.44 20.90
Forest Meadows:
Afralee - Adnasad 1000 4.88 15.92
Khandooshan 500 0 3.48
Bozaghan 1000 4.88 20.90
Takhté - Korda 1500 7.32 17.91
Transition Zone:
Tunnell Solaiman Koshté 1500 2.44 16.94
Upper Karkoly Valley 2000 60.97 0
TOTAL -- 100.00 100.00

 

42

Seasonal Food Availability

The largest average supply of available herbaceous forages
was recorded in summer, when it reached as much as 1253.4 kg dry
wt/ha (Table 18). This was in contrast to the winter supply of
only 11.5 kg dry wt/ha on the steppe.

0n the average for all vegetation types, the largest supply
of browse also was recorded in summer (Table 19). A drastic volume
reduction in winter was due to the loss of leaves and to the dis-
appearance of bushes under snow. The highest share of browse
forage was produced by oaks (Table 20).

Ocular estimates of the acorn crop were calculated in kg dry
wt/ha. It is recognized that approximate methods were used in this
regard but it was felt that even a rough estimate was better than
none at all. It was found in the Caspian deciduous forest 38.7
percent of the largest oak trees produced an acorn crop of 156.4 kg
dry wt/ha. The crop produced in the transition zone was estimated
to be 119.6 kg dry wt/ha and 42 percent of the trees produced acorns.
Acorns possibly are the major fattening feed in the fall for maral

as well as for other wild life, especially wild pig.

Food Use and Preference Values

 

Food composition determined by observations of wild maral

were not amenable to precise quantification. Yet it is believed

43

TABLE l8.--Seasonal changes in the average available herbaceous
forage supply in the Mohammad Reza Shah National Park,
Iran, 1976-1978. All values are given in kg dry wt./ha.

 

 

 

Vegetation Type Spring Summer Fall Winter*
Caspian deciduous forest** 54.6 264.8 292.4 28.3
Transition zone 153.6 1253.4 743.3 35.1
Steppe 126.8 432.6 312.9 11.5
TOTALS*** 108.3 503.6 354.6 19.1

 

*Available during snow-free days or mild winters like
winter of 1978. Totally absent for about 2 months in winter 1977.

**Including forest meadows.

***
Weighted according to the areas involved.

44

TABLE 19.--Seasonal abundances of the browse in the Mohammad Reza
Shah National Park, Iran, 1976-1978.

given in kg dry wt./ha.

All values are

 

 

 

 

 

 

Vegetation Type Spring Summer Fall Winter
Caspian deciduous forest* 51.16 415.55 153.48 29.16
Transition zone 40.53 212.84 94.62 27.72
Steppe 13.76 29.44 24.44 7.78

TOTALS** 27.86 164.73 70.64 19.33

 

*Including forest meadows.

*1:
Weighted according to the areas involved.

45

TABLE 20.--Browse supply (kg/ha) by species in the Mohammad Reza

Shah National Park, October 1977.

 

Caspian

 

 

 

 

Species Deciduous Tragzzzion Steppe
Forest
Cotoneaster sp. 1.09 3.45 0
Lonicera floribunda .75 1.90 0
Rhamnus sp. .96 3.39 0
Euonymus sp. 1.95 7.86 0
Colutea persica 2.79 11.32 1.14
Ephedra sp. 0 5.09 22.95
Acer ibericum 0 68.30 3.01
Carpinus betulus 56.84 0 0
Viscum a1bum* 3.96 0 O
Quercus sp. 301.10 93.46 0
Acorns 156.40 119.60 0
Other 50.09 18.07 2.34
TOTAL** 415.55 212.84 29.44

 

*Viscum is a saprophyte. A potential winter food in harsh
winters, it grows mainly on Carpinus branches.

**
Acorns and Viscum are not included.

46

that some insights into the feeding habits of the maral were
gained.
In winter season, Colutea persica, Euonymus sp., and Quercus

castanaefolia were preferred browses (Table 21). In spring,

 

Euonymus sp., Ephedra sp., and Acer ibericum were highly preferred

 

Table 22). Capsella bursa-pastoris, Phleum paniculatum, Poa bulbosa,

 

Vicia persica, Dactylis glomerata, Onobrychis gobi, and Coronilla

 

varia were also among the preferred forage species in that season

(Tables 23 and 24). In summer Acer ibericum, and Quercus

 

castanaefolia were highly preferred (Table 25). Dactylis glomerata,

 

 

and Festuca ovina were the preferred forage species (Table 25).

 

And finally in the fall, Dactylis glomerata, Poa bulbosa, plus

 

acorns and Colutea persica were the preferred foods (Table 26).

 

Agropyron trichophorum, and Festuca ovina were the most

 

 

abundant herbaceous species year around. In terms of bulk contribu-
tion to the maral's diet, the most important forage species was
(Agropyron trichophorum, a grass (Tabls 23). 0f the 19 plants eaten

by maral, Dactylis glomerata seemed to be the most preferred forage

 

species on an all-year basis (Tables 21, 22, 23, 24, 25, and 26).
The oaks were the most abundant food sources in the park. They
produced the highest amounts of fruits as well as of browse, and in
terms of bulk contribution to the maral's diet, they were the most
important browse species.

The Persian red deer is mainly a grazer, preferring grasses

over woody forages.

47

 

 

 

 

mo.om oo.oo_ oo.oop Fo.w mm.mm 4<~0h
o o o wn.__ o p~.e gmgpo
mo. mm. Fm. mm.op eo. p~.¢ mumwgguumcmam magampmm
no. cm., mm. Fm.“ 0o. Np.m .am mammmuwgu
m_. oc.~ RN. Pp.m No. em. .am 0::0gau
~_. ¢¢.m mm. om.¢ mo. ow.~ mzpzuwa mzcwagmu
mm. Pm.¢ we. wo.~ co. mm. snap: Esumw>
om. em.“ Fm. mm.P mo. mm. .mm gmummmcopou
we. Fm.w mm. em.~ mo. om. accanmcopw 0:00mcog
mm. em.o_ om.~ mm.m 4N. m~.~ aawamau macs»:
em. 05.0P mu.~ mn.m NN. m¢.P 530mg0aw :00<
om. mp.mp Nm.p ne.p pp. mm. .am macsmsm
mm. m~.- _—.mm mm.m~ mo.~ 0m.m mwpommmcmummu msongo
em.p Fm.om m_.op om.w mm.” om.m .qm masx:o:m
mm.~ m~.~m m¢.mm no.mp mp.m ~¢.m muwmgwa mmpapou
mmcwwmm 00>020m 00w: 0—nmpwm>< 00>050m 0anpmm>< managed Auooz
00:0:000LQ mmmmucmugma ~E\.o: mango:

 

.Nump Lapewz .xtaa Facowpaz
cmcm m~0m 00:50:02 .mpcsou 00*: x: 00cwsgmpmu mm mmcwpm: 00:0:000:a 0000 —0:mzuu.- m4m<p

48

 

 

 

 

oo.oo_ oo.oo_ m~._ 00.: 0:000
00.0 0.0 _0.0 0N.m Fe. 00. .am 03:50::
00.0 0.0: 00.0 00.0 00. mm. 000000: 0003000
00.0 ¢.¢_ 00.00 00.00 00. 00.0 awpommmempmao maggwao
N_._ 0.0, 00.00 00.0N ma. 00.0 2:00:00: 000<
00., 0.00 :0. 00. Po. mo. .am acumgam
mN._ m._N mm.om 40.00 mm. 0_.F .am mzexcosm
00>0E0m “00¢ 0Fn0pw0>< 00>0E0m 0Fn0pw0><

mmcwu0m

00:0:00000 m0m0u:00:00 0:\mx 0000000 acooz

0:000: xgc 000000

 

.mnmp 0:30 .x000 F0cowu0z :0cm 0~0m 00:50:02 .0:=o:m 0:00000 mchQm apoxg0¥
00 00: z0uuepp 0 0:0000 000:0 0P0: N 0:0 mama 0P0: 0F .p0g0s _m an :0000530:00 0mzogmuu.mm u00<~

49

 

 

 

 

 

 

 

00 00 00.000 00.000 00.000 00.000 00000
- - - 00.0 - 00.0 00000
00.0 00.0 00. 00.0 00.0 00.00 00000000000 0000000:
00.0 00.0 00.0 00.0 00.0 00.0 0000000000 0000000:
00.0 00.00 00.0 00.0 00.0 00.00 00000000000 000000
00.0 00.0. 00.0 00.0 00.0 00.00 000000000 000000
00.0 00.00 00.0 00.0 00.0 00.00 00000000000 000000
00.0 00.00 00.0 00.00 00.00 00.00 000>0 0000000
00.0 00.00 00.0 00.0 00.0 00.00 00000000 000000
00.0 00.00 00.0 00.0 00.0 00.00 0000000000000 000000000
00.0 00.00 00.00 00.00 00.00 00.00 000000000000 000000000
00.0 00.00 00.0 00.0 00.0 00.00 00000000 000.0000
00.0 00.00 00.0 00.0 00.0 00.00 0>0000 000>0
00.0 00.00 00.0 00.0 00.0 00.0 00000000 000000:
00.0 00.00 00.00 00.0 00.00 00.00 0000 00000000000
00.0 00.00 00.0 00.0 00.0 00.00 0000) 0.0000000
00.0 00.00 00.0 00.0 00.0 00.0 0000000 00000
00.0 00.00 00.0 00.0 00.0 00.00 000000000 00000000
00.0 00.00 00.0 00.0 00.0 00.00 0000000 000
00.0 00.00 00.0 00.0 00.0 00.0 00000000000 000000
00.0 00.00 00. 00. 00. 00. 00000000-00000 00.00000
00>0000 0000 0000000>< 00>0000 0000000>0
0000000 0000000 000000
00:0:00000 00m0uc00000 0:000 .

000003 000 000000

 

.0000 0:00 .0000 00:00002 :00m 0000 0055000: .0000 0:00000 0:0:0m 0000:0x 00 00: 000-000 0
0:0000 00000 0003 0 0:0 .0000 000: 00 .00005 00 00 00>0s00 0000000 000000000; 0:000zuxgonu.mw mam<0

50

 

 

 

 

 

 

 

00.00 oo.oo_ oo.o00 00.0 0.00 0<0o0
. u n 00.0 n 0.0 0000000 00:0o
om. 0o.m_ 00.0 00.00 m.o 0.0 .00 0:00:00“
00. 00.00 00.00 mo.0~ 0.0 0.00 0000000 0000000
00.0 00.00 «0.00 00.00 0.0 0.00 0000000:0000u 0000000
00.0 00.00 00.00 0o.N~ 0.0 0.00 00000000 0mu<
"mummh oz< mmamzm

00.0 oo.o00 oo.oop 00.0 00.00 0<000
. u . 00.00 - 00.00 000000m 00:00
00.0 00.0 00.0 00.0 mo. 00.0 000000000: 0000000:
00.0 00.0 00.0 00.0 00. 00.0 0>0u00 0:m><
00.0 mo.op 00.0 00.0 NP. 00.0 0000000 000
00.00 00.00 00.00 00.0 00. 00.0 000000000 00000000
"mmmou oz< mm<mu

00>osmm 000: 0000000>< um>oemm 0000000><
wwwmwwmw0 0000000 000000
0 00000000000 ~e\.o: 000000

 

.0000 000000 .0000 00000002
:00m 0000 00550002 .:0000>0mmno x: 00:05:0000 00 0000000 00:0000000 boom 0000:--.0N u00<0

51

 

 

 

 

 

 

 

mp.o oo.oop oo.oo~ Fm.~ mm.m~ 4<p0h
. u . Po.¢— - pp.e gmgpo
mm. em.~ mm.m mo.¢m up. cw.“ muwmgma mmuzpou
NF.~ mm.o mm.¢¢ mo.o¢ Pm. mn._F ampommmcwummu mzugmzo
mp.~ w~.mp mm.m¢ m~.P~ mm. m~.o Ezuwgmam Lmu<
”mummb oz< mmzmzm
em.m oo.oo~ oo.oop ¢.m~ m.oom 4<~0h
. u . ¢N.uw . ~.-m gmgpo
mo.m mm.~m mp.np mm.~ o.m m.m amonpza mom
Fm.“ up.mm ¢~.mp so.~ ¢.e m.u Ezwpomwumsmcm E:P_omwgh
Fm.“ -.am em.- No.~ ¢.m m.op m>wumm acm><
mm.“ m~.mm N_.m~ Fm.m w.o ~.o_ wcw>o auspmom
mm.m mo.m~ ~o.- m~.~ o.m e.m mumngoFm mwpxpumo
"mmmom oz< mm<mo
um>oemm umwo mpnmpwm>< um>osmm wpnmpww><
wwwmwwmwg wwwuwam mango;
$ a mmmmpcmogma ~E\.oc mango;

 

.N~m_ gwsssm .xgma chowawz zmgm

mNmm umssmcoz ._mgme u_w3 mo :owpm>gwmao an vmcwELmuwu mm mmcwumg wucmgmkwga woo» ngmzun.mw m4m<p

52

 

 

 

 

 

 

 

om.¢P oo.oo~ oo.oo_ _F.m mm._~ 4<FOP
o .I o o ap.~ 0 Ne. gmguo
Km.o mm.w -.~_ ~N._~ mm. mm.¢ asngmnw gmuq
no._ mm.m_ o~.mm mm.¢e No. Ne.o_ cypowmm=mpmmu maugwac
No.P No.mp mm.- NN.p~ _~. mm.¢ mcgou<
mo.~ me.om mN.PP mm.m mm. mp.. mummgma mmp=.ou
”mummh oz< mmamzm
_~.mm oo.oop oo.oop Fm.m_ mm.me 4<h¢k
o o o oo.om o ¢F.w_ gmgpo
«F.P _m.mm No.0, mm.ep Pm.m ofi.m a=w>o wuapmmm
N_.P mm.¢m op.o NN.m mp.p m¢.m m>wwmm m=m><
Nm.F om.mm ou.m mo.m _N._ mm. E=V_O$wpm=m=m s=w_omwgp
so. em.m¢ mm.kp mm.op m¢.m mm.o 23L0pump mzsogm
-.F eo.Fm mo.- mo.mp ¢m.m om.op Eagogaoguwgp cogxgogmq
No.~ Pp.om um.o_ Ne.m ¢F.~ om.m wwgomsmp mao_me<
mm.~ $0.00 mm.m .m.~ mo.P mm._ amon_=n no;
mm.~ mu.o~ om.m N©.N m~._ mN._ mumgmso_m mPPAgumo
"mmmom oz< mm<¢w
ma=_pm¢ um>o§mm “ova m_nwp_m>< ww>o§ma wpnmpwm><
mocmgmwmga mmmuwam mamgon
mmmmucmugma ~E\.oc mmmgou

 

.nmmp Ppmm .xgmm
chowumz zmsm mNmm umssmcoz .cowum>gmmao an cmcm5Lmumu mm mmcwumg mucmgmmmga coo; pagmzuu.o~ m4m<h

53

The amount of browse utilized by an average maral was esti-
mated to be 2.87 kg/day (2.8% of body weight) during the winter of
1977 (Table 27). This seems to be normal when compared with food
consumption of American elk (Telfer and Scotter, l975). The amount
of forage (browse, grass, and forbs) utilized by an average maral
was estimated to be 3.6 kg/day (4.3% of body weight) during the
spring of 1977 (Table 28).

Competition for Food

 

In addition to maral, other large herbivores also present
in the park were wild pigs, goitered gazelles, wild sheep, wild
goats, and roe deer. An attempt was made to estimate their numbers
in the park (Table 29). The extent to which the recorded diet of
maral was overlapped by those of the other five ungulate species
was calculated (Tables 30 and 31). The competition coefficient (cc)
values were determined to be .16, .08, .06, and .02 for wild pig,
wild sheep, roe deer, and wild goat, respectively. The gazelles with
a habitat totally separated from maral, had a cc = 0. Wild pig and

maral competed heavily for acorns in the fall (Table 30).

Range Condition and Trend

 

More than 8 percent of the available shrubs were severely
hedged in the transition zone by browsing deer (Table 32). A further
fraying (debarking by rubbing antlers), threshing, and rubbing of
sapling trunks was caused mainly by male deer, using their antlers.
Stripping was done by both sexes using the teeth to cut through the

bark and to tear off strips by an upward movement of the head. In

54

.pgmwoz o>wp m.poew:o on» oo acoogmo m.~ ooposoo

xoo Loo oo23mcoo ooow ogu .omogo>o oz» co .mx mm.mo u Looo m— x ax nw.~ mo: ocoogm mcwgoucwz on»
um oomuopoooo Poona on» mo :owuo23mcoo woo» Pouop .mx mmm_ oocmwoz Focus mp asp cosy .coom ogo;

Foams ago co »_aaa .gaau no; gmopoa Loo Acumpv mezo_owoao~o an omgmwpnza moan “comm: soon «no mo

ammo flopo

 

 

 

 

 

 

ox mm.~ u No;\mx oo.ov an; ommv u zoo coo Pages omogo>o no mo :opoo monogoo ugmwoz->go
flow: mo compogaov AcowpoF=ooo Fogoev
osxmx o; u moo; cowposomcoo wood
Acowuosomcoo omocomv Aocsogm momgmucwz mo owcov
mm.~ oo.oop oo.oop oo.o mm.nn_ 4<hop
o o o mp._F o Pm.mp womwgnouocwom mogswpoo
m~.o mo.o oo.m_ m~.oo mm.o om.m~ owpowoocoumoo moogozo
mo.o Po._ mu.o so._ mo.o nm.~ .om mgoogom
nn._ um.m m~.op mo.m um.o om.o~ oowmcoo monopoo
oP.N _N.o oo.- mm.om mm.~ so.po .om moszcozm
oo>osoa “owe opoopwm>< oo>o2om opoopwo><
mmcwuom o
mococommgo monoucoocoo o;\mx mowoo m omzocm

“sumo: ago omngou

 

.unmp xgmogoou oco monsoon .xgoo pocomuoz zoom o~om ooEEocoz .ocoogm mcvgoucwz wpsmox
coswopom no oowgoo monumm o mcwgoo Amoum moonwalzppow p oco .mocw; mesons o .mocw;
mcoox m .mmoum oxvom N .mmcppgoox opoeom my Focus op an couoo momogom pcmwmz-»gon-.nm mom<p

55

TABLE 28.--Food consumption rate by 31 maral during a 114-day spring
use at Karkoly Spring, Mohammad Reza Shah National Park,
Iran, June 1977.

 

ANIMAL POPULATION:

Food habit studies of the maral were carried out at Karkoly-Spring
where approximately 300 hectares supported a population of 31 maral,
16 wild pigs and 2 wild sheep. Spring and summer observations
showed that the marals used this site daily while pigs and sheep
were less frequent visitors. During an ll-day survey, maral herds
were seen daily as compared with 2times for pigs and once for sheep.

 

31 maral + 16 wild pigs + 2 wild sheep= 49
Average Adult Weight:

Red Deer = 168 kg (Dzieciolowski, 1970)
Wild Pig = 150 kg (Van Den Brink, 1968)
Wild Sheep = 60 kg (Firouz et a1., 1970)
or in animal units: Nild Sheep = 1.0
Wild Pig = 2. 5
Maral = 2. 8
Total Animal Population = Sheep. 2 E1.) .0)
Pigs. 16 2. 5)
2. 3

Mara1:]31.g 2+
n1ma Units

Since energy intake is proportional to energy expended in metabolism
(Blaxter, 1962), total forage removed was approtioned to each species
population according to the energy metabolized by each population.

If all 3 game species are equally competitive for food, then

the maral share of food removal was 3}462é8 x 100 = 59.1%

Maral daily forage consumption rate =
(300 ha) (115.34 kg/ha) (59.1) = 3 6 k
49 an ma 4 days 00 ' g
Maral population consisted of:
10 calves, 3 young hinds, 2 spike stags, 9 young but fertile
hinds, 2 mature stags, and 5 mature and old hinds.

The population weighed approximately 2582 kg. Average food
consumption was estimated to be 4.3 percent of live weight.

 

56

TABLE 29.--Estimated numbers of major prey and predator species in
Shah National Park, Iran, 1976-1978.

the Mohammad Reza

 

Animal Species

Total Population in the Park

 

Preyg§pecies:

 

 

 

Wild Sheep] 10,000 - 11,000
Wild Goat2 4,000 - 4,500
Wild Pig3 2,500 - 3,000
Maral3 1,900 - 2,100
Roe Deer4 500 - 700
Gazelles 250 - 300
Hare6 8,000 - 12,000
Pheasants6 1,500 - 1,600
Chukar6 3,500 - 5,000
Predator Species:6
Leopard l6 - 22
Wolf 22 — 29
Cheetah 3
Bear 22 - 24
Jackals 50
Fox 60
Wild Cats 100
lBased on actual counts of animals seen along transect
lines established for sheep.
2Based on full day observation in sample areas.
3Based on actual counts of animals seen along transect
lines established for maral.
4Based on full day observation at wintering grounds in
January and February 1977.
:Based on herd counts at Mirza Bailou Plain.

My impression of animal populations based on limited

information of animal signs and few sample counts.

57

 

 

 

oo.oo_ oo.oop oo.oop oo.oo. oo.oo. oo.oop o<»o»
1mm. as Se in mm. MM is
o o ~.m m.mm o o mccoo<
o m.m~ a.“ o ~.o o .am maocaao
o o e.“ o o.o _.o 2:0»zma» zmu<
N.o o m.o o m.oF m.m .am a»m»saoc<
_.m o «.4 o ¢.¢_ a.» .am azomgau
m.NP _.m P.m~ m.ep N.m ¢.o_ momeanpm m»»»oumo
¢.Nm w.“ m.N_ m.op N._N ¢.om a=»>o aosommo
xm.m sm.m $5.5 &_.m so.m &_.m_ amon_=a mom
3...“. a... a... m...” ..._..,...

 

.mnmpummmp

.coLH .xgoo Poco»uoz smgm oNom oosaogoz .mo»ooom mango» m »o oom»gosoo mogo>»ocog
omen» so» op=:»s Loo mop»o »o songs: on» so oomoo um»o oo>gomoo ogp »o moo»p»ooosou-.om mom<p

58

TABLE 31.--Competition for food between maral and 5 other large
herbivores, Mohammad Reza Shah National Park, Iran,

 

 

1976-1978.
A B A x B/10000
Species Diet Range Competition
Overlap Overlap Coefficient
% %
Wild pig 34.2 48.2 .16
Wild sheep 35.4 22.4 .08
Roe deer 21.7 30.2 .06
Wild goat 34.2 5.4 .02

Gazelle 36.3 0 0

 

59

TABLE 32.--0ccurrence of degrees of hedging on four major browse
species, Mohammad Reza Shah National Park, Iran,

 

 

 

1976-1978.
Plant Species Eggfigggd Hedged form (in percent)*
0 l 2 3** 4**
Colutea persica 125 49 22 16 13 0
Euonymus sp. 115 54 22 16 8 0
Rhamnus sp. 44 55 45 0 0 0
Acer ibericum 34 60 40 0 0 0

 

*Based on Riney et al. (1959).

**Severely hedged = 1;); I “95 I 42 i 32 = 3%— = .08 = 8%

 

60

the park, maral were seen to use the bark of Euonymus sp., Prunus

caspica, Juniper sp., Rhamnus sp., Colutea persica, Carpinus sp.,

 

 

Acer sp., and Quercus sp. The debarking (fraying) of trees, however,

is not yet a serious matter in the park (Table 33).

Carrying Capacity

 

The range carrying capacity is defined as the maximum number
of physically fit maral that can, in the park ecosystem, survive
through the least favourable environmental conditions within one
year.

Hedging and debarking of shrubs and trees in the transition
zone indicated that maral populations there had reached the range
carrying capacity. Maral consumed 2.87 kg of browse per day there
during winter (Table 27). There was 27.72 kg of winter browse
available per hectare in the transition zone (Table 19). For a
period of three months with 60 percent allowable utilization (Telfer
and Scotter, 1975), the transition zone of approximately 14,000
hectares could support a population density of one deer per 15.6
hectares or 900 maral. The best estimate of maral population in the
zone, based on the fall pellet-group counts (Table 7), however, was
1064 maral, 164 maral were in excess.

While the maral is the major and most abundant ungulate in
the transition zone, it is outnumbered in the Caspian deciduous forest
by both wild pigs and wild goats. Based on animal signs and several
population censuses, there were approximately 3000 wild pigs, 700

roe deer, 1200 wild goats, and 830 maral. without further study, it

61

 

 

so, go Nom 1 oaoasmo mo=a_a o<»o»
mmm_ u om»_Pao mo=a_a o<»o»
om ~_ mm mm» om_ me so anaap zao<
om op om Nm_ o_N ON mm m=_=oma m==»azmo
mg m_ cm mom omm ow mm m»»o»am=aommo msuzmso
o“ o mm mm_ omm o“ m“, mu»mcaa amoapoo
swam so so
oo>osom omxogm
o»E»o o»ewo o=a_a cussed
mococo»soog»o »o cooszz oagsm Lo mos» ooaogu
»o “cmocoo LouoEo»o Lozoo goon: »o p:m»o:
omoco><

 

.mmmpuonmp .cogH

.xgoo Poco»poz gogm oNom omsEogoz .Fogos »o co»>o:oo m:»zoguuu.mm m4m<H

62

is difficult to determine the range carrying capacity of the

Caspian deciduous forest for maral. But considering maral preference
for forest meadows in winter (Tables 14 and 15) and 39.16 kg/ha winter
supply of browse, meadows can support a population density of one

deer per 11.0 hectares. The best estimate of maral populatin

density in the zone, however, based on the pellet-group counts

(Table 7) was 29.3 ha/deer. It was thought that some other limiting
factor, other than available winter browse, caused a lower carrying
capacty of the range there for maral. Probably this was competition

by the pigs.

Physical Condition of Maral
In the U.S.A. an inadequate food supply for deer is commonly
observed (Gwynn, 1965) to produce poor quality antlers. This has
also been noted in European deer (Lowe, 1971 and Whitehead, 1972).
According to Gwynn (1965) deer herds in the very best ranges average
about 5 percent spikes (i.e., yearling had antlers with 2 to 6
points, spikes here referred to the yearling stags with a poor set

of antlers; 2-pointed). The observed average of 20.1 percent

31 spike
2-6 point

 

spikes (154 ), then, might be an indication of the maral

population being in poor condition.
Of 975 animals classified at the end of the summer period
when they were expected to be most fit (Mautz, 1978), 11 percent

were judged to be in poor condition (Table 34).

63

TABLE 34.--Physica1 condition of maral in the Mohammad Reza Shah
National Park, Iran, September 1977.

 

Number

Body Condition

 

 

Vegetation Type Examined Good Poor
% No. % No.
Caspian deciduous forest 430 94 404 6 26
Transition zone 545 85 463 15 82
TOTAL 975 89 867 11 108

 

64

Mortality Causes

Of the 27 kills found, only 11 were of maral (6 males and
5 females). Five of these were killed by leopards, two by wolves,
one by a car, and three by poachers. Leopards and wolves were the
major predators capable of preying on maral in the park but, from
information provided by the villagers, local poachers killed more
adult maral than park reports suggest. Though based on limited data,
the maral was the most preferred prey species for leopards and the
second most important prey species for wolves (Table 35).

If it can be assumed that a leopard consumes an average of
1000 kg of meat per year (Schaller, 1972) and that on the average
25 percent of a kill consists of inedible portions, then where
27.8 percent of this represents maral it must be that three maral
are taken annually by each leopard. Pimlott et a1. (1967) estimated
the average daily meat consumption of a Canadian wolf to be 4.5 kg.
Assuming that on the average, 25 percent of a kill consists of
inedible portions, each wolf has to kill 2010 kg per year to survive.
This would indicate that the wolves in the park, whose food con-
sisted of about 77.8 percent prey of medium size and 22.2 prey of
larger size (Table 35), each kill about three maral. The number of
wolves and leopards in the park are believed to be 29 and 22,
respectively (Table 29). If these estimates are accurate, then
these two predator species kill about 153 maral annually, an
estimated kill level of 7.7 percent. According to Mech (1970) and
Schaller (1972, 1977), predators may become the major controlling

mortality factor where prey-predator ratios fall to 100-150 prey

.ooop .zamem»a new mauzzoaa :0 oamam "momaom

 

 

65

 

 

oo.oo_ oo.oo~ oo.oop mm 4<»o»
11 11 11 11 m.m 1 some mom
w.» 11 F.P_ 11 m.» P oPPoNow
11 no.» 11 m.mm o.o~ o poow up»:
FF.» mm. m.mm ».o_ o.om m goosm o_»2
on. mo.» P._F N.NN o.m— m m»o o_»3
NN.N w».~ ~.NN m.»~ o.op m Pogo:

mo>po3 mogooooo FP»¥ ppmx “moocoo mFP»¥ o x
kmmc»uom oocmco»oco =»mmnwwaoo cwonmowwwm mpowflwo>< coosoz mo»oo m moo

 

_ .mPF»x oo>comoo »o goosoc Fposm o co comma amnmp1mnmp
.cogH .xcoo pogo»uoz gogm onx ooEEogoz mgu :» mm>~oz oco mogoooop »o moocogo»ogo zoso11.mm u4m<p

66

animals per predator or less. At higher ratios, however, predation
becomes only one of several other contributing mortality factors and
cannot normally be considered to be a primary controlling influence.
Since there were 256 prey animals per predator (Table 29) in the
park, it seems that the kill rate is too low for predators to act
as the regulatory or controlling factor.

Red deer herds reject diseased animals in their midst
(de Nahlik, 1974). Lone hinds often are (calving season apart) sick
specimens and according to European standards (de Nahlik, 1974)
should be shot. Winter observations showed 124 lone hinds (Table 8),
possibly sick animals. No information is available on diseases
carried by maral in the park but in Europe, the most common ones are
louping-ill and tick-borne fever or rickettsial disease (Whitehead,
1950). Parasitic worms were the major cause of mortality among

calves during the late winter and spring in Europe (Whitehead, 1950).

DISCUSSION AND RECOMMENDATION

Though deer are known to be highly selective in the food
they eat, their distribution is not usually governed solely by the
occurrence of a particular food species (Staines, 1974). Varying
regionally in their occurrence or in their selection by deer, pine
was an important food item in Dutch and Danish studies during late
winter (Jensen, 1968), it was unavailable in the Scottish one
(Staines, 1970), and was regarded as starvation food in Poland
(Dzieciolowski, 1969). The highly selective nature of deer for food
does not imply that they occupy areas only where a particular food
species occurs, it is merely true that deer select from what is
available. The presence or absence of nutrient chemicals in the
soil also affects the degree to which deer are attracted to food
plants (Staines, 1974). Food affects deer distribution daily and
seasonally. Staines (1974) suggests that though the quality and
quantity of food may be important in the distribution of red deer in
spring and summer, it is probably not the main limiting factor on
winter ranges. The relationship between food and availability of
shelter also is important. Where winters are severe, black-tailed
deer are seldom found feeding more than 100 yards from shelter
(Taber and Dasmann, 1958).

Meadows are essential to maral in the woodland and their

maintenance is of prime importance. Though the preservation of

67

68

natural conditions is the primary objective in national park manage-
ment, the maintenance of at least some important meadows perhaps
should be given priority if tourism is affected by maral abundance.

The vegetational evidence suggests that the maral range is
changing in the direction of a replacement of preferred maral foods
by undesirable forage species. An invasion by woody plant species
has now progressed to an advanced stage and to the point where
mechanical or chemical treatment must be applied in several areas.
Since control by these methods results in dramatic changes in the
landscape and there is now much concern about the environment, high
priority must be placed on developing techniques and procedures which
are consistent with national park objectives and which nevertheless
meet management needs. The preservation of natural beauty must be
a primary objective in plant control project planning. Plant control
treatments must be aesthetically pleasing.

The meadows in the park once were cultivated plots. These
areas, now about 20 years old, currently are the primary feeding
sites for maral. They are also vital to other species of wildlife,
particularly the more than 1500 pheasants in the park. What then
are guidelines for meadow maintenance projects?

The management of animal life in national parks is so
closely associated with the management of the vegetation which
supports it. Forest meadows in the park are successional communities.
It is, therefore, necessary to manage the habitat to achieve or

stabilize a desired stage. 0f the various methods of manipulating

69

vegetation, the controlled use of fire is the most natural to apply.
0n Isle Royale, in Michigan, USA, moose range is created by periodic
blazes that open the forest canopy.

In comparison with American and European findings (Harper
et a1., 1967; Lowe, 1969; and de Nahlik, 1974), the Qgrvu§_population
represents a high standing crop of low productivity. Park maral are
growing and reproducing well below their potential rates.

The condition of the maral, coupled with the low calf:hind
ratio, suggests that the population at the time of the study was above
its range carrying capacity in the important transition zone of the
park. This may make the population particularly sensitive to the
stress of severe winters. Overstocking and range depletion give an
advantage to females which have lower food requirements than males
(Darling, 1939; Anderson, 1958; Flook, 1970), and results in a dis-
proportionate sex ratio (Cowan, 1950; Taber and Dasmann, 1954;
Robinette et al., 1957; Klein and Olson, 1960; Peek et a1., 1967).
Heavy selective shooting in the park also may have contributed to
the disproportionate sex ratio. Both overgrazing and overshooting
can be expected to cause a loss of high-quality stags in the park.

Stags outnumbered hinds in winter at the higher elevations.
Although separation of the sexes helps limit intra-specific compe-
tition for food (Geist and Petocz, 1977) winter mortality will be
higher for stags which winter where the snow is deeper and the
temperature colder than on the lower ranges. The existence of
appreciable numbers of mature yeld hinds (females that did not

reproduce during the previous season, though not necessarily barren)

70

in the park population is symptomatic of a nutritionally inadequate
environment, as also recorded for American elk (Greer and Howe,

1964; Harper et al., 1967). The low breeding success of hinds in the
park can be explained largely in terms of the slow rate of recovery
from the effects of pregnancy and lactation as a result of poor
forage. This in turn affects the ovulation and conception rates.

According to Mitchell (1976), aspects of reproduction in both
sexes of adults also are influenced by body weight and condition.

For stags, the spring season is the most important since extra
nourishment is required for antler growth and to regain physical
condition.

Next in importance is the period following the rut, when the
main need is albumen and carbohydrates to provide flesh to cover
bones for as much as 14-17% weight has been lost during this period
(Mitchell, 1976). The hinds require about half the quantity albumen,
calcium, and phosphoric acid (for reproduction and lactation) needed
by a strong stag during the eight-month period October to June
(Whitehead, 1950). The size of antlers have been related to food
supply, soil, light, and body size (Chapman, 1975).

Our knowledge of the genetics of deer is very limited and any
attempt to alter antler characteristics by breeding, which often means
selective shooting, rests largely on chance. Cadman (1967), for
example, states that the antler-quality of fallow deer is controlled
by shooting the less-desirable males selectively. While antlers
are only carried by the male maral, their structure is affected by

the genetic characteristics of both sexes. Selective shooting of roe

71

deer in Denmark did not increase the size of the antlers. Improved
antlers were only obtained when the entire herd was killed and
replaced by deer from another area where the deer grew larger
antlers (Anderson, 1961). The trophy value of red deer antlers in
many areas of Scotland has declined during this century despite
efforts to stop or reverse it (Lockie et al., 1967; Lowe, 1971).
There are experiments carried out in Britain today, however, which
prove that with dietetic additions it is possible for randomly
selected hill calves to develop six to eight pointer on its first
rack at the age of about 10 months (de Nahlik, 1974).

A reduction in maral density in the park certainly would be
reflected in improved reproductive and growth performances of indi-
vidual animals. This could best be accomplished by removing the
excess animals, probably more than 160. Perhaps the most feasible
procedure is by controlled shooting but to avoid the encouragement
of unregulated poaching and to insure the selective greater removal
of reproductive age hinds and poorlv-antlered stags, onlv park
personnel should be involved in the hunt.

The adjustment of herd size may have to be carried out over
a period of several years. There are a variety of requirements to
be met in a well-devised and well-executed shooting plan:

1. The maintenance of a static-size population, adjusted
to the carrying capacity of land with a correct ratio of sexes.

2. Adjusting deer numbers so as to achieve a density of one
maral to 15.6 hectares (in the transition zone) as the highest density

acceptable.

72

3. The achievement of a correct number and correct age
structure.

4. The elimination of poor-quality old animals.

When confronted by a herd, it can be difficult for the park
official to make a quick selection of a suitable animal to shoot.
Certain points of guidance can be followed:

1. The leading animal may not be shot, unless it is necessary
to split the herd. It is especially important when hind groupings
are small, that the leader (normally a dominant female) should never
be shot. But conversely, if hind herds are larger than desired, the
shooting of the leading hind may result in a desired split-up of the
herd.

2. Lone hinds normally (calving season apart) are sick
animals and should be shot.

3. Each female without a calf should not automatically be
shot, it is fairly certain that after a 'rest' such a hind can
produce a strong calf.

When it is desired to keep numbers static:

l. The total number of animals to be shot should equal the
net annual increase.

2. Stags of poor quality should be eliminated, ideally in
their first autumn. A quick rule is that the antler in the first
head must be at least equal in length to the ears. An antler which
can be inscribed into a rectangle is one with a good future, whereas

a triangular pattern is unlikely to develop well (de Nahlik, 1974).

73

Red deer, espeically hinds, are very fond of chewing cast
antlers or any bones they may find (Whitehead, 1950) in the late
Spring and early summer. This is the antler growing season for the
stags and calf-rearing season for the hinds. The collection of
antlers by the park personnel should not be allowed since they provide
appreciable amounts of calcium (28%) and phosphorus (22%) during the
critical spring season (de Nahlik, 1974). Yet, according to park
personnel, an average of 600 kg of good quality antlers are collected
annually. This comprises 25-30% of all shed antlers and about 80% of
all the high-quality antlers.

In considering a program to insure maintenance of the park
ecosystems, it is suggested that:

1. Assessments of animal populations, range interactions and
dynamics should be conducted annually in all important areas of the
park and for all major animal species. Basic information is essential
for the maintenance of both the park's animal inhabitants and its
habitats.

2. Particular efforts should be made to identify areas where
range use conflicts may occur between the maral and other species.
Any ungulate range interactions which seem likely to endanger the
quality or quantity of habitat conditions should be managed in favor
of the restoration of natural conditions.

3. Several exclosures should be built, each perhaps 0.5
ha in area. These would exclude all large grazing animals so that
comparisons could be made between grazed and ungrazed vegetation.

Such exclosures would enable early detection of range over use.

74

4. Study of the effects on the range of rodents and hares and
other small mammals is desirable in order to determine their importance
as influents of environmental conditions. Smaller exclosures should
be built to exclude small mammals. These may be made as a part of the
larger exclosures.

5. In order to determine the range carrying capacity more
accurately, seasonal food and nutrition requirements should be deter-
mined. Chemical composition of forages should be studied. As pointed
out by Mautz (1978); the analysis of nutritive values of white-tailed
deer foods perhaps should be shifted from an emphasis on winter forages
to studies of summer and fall food availability and digestibility since
these determine the accumulation of body fat. Browse evaluation
studies have typically shown that most natural winter browse foods are
poorly digested by white-tailed deer (Ullrey, et. al., 1964, 1967;
Robbin and Moen, 1975; Mautz et al., 1976; Mautz, 1978). It now
appears that the storage of a large energy reserve in the form of fat,
prior to the winter season, is the major factor determining the sur-
vival of northern white-tailed deer (Mautz, 1978).

6. Regular surveys to appraise the occurence of erosion,
depletion and pollution should be made with respect to park preser-
vation.

7. In any case, where management is undertaken in order to
preserve wilderness values, periodic assessments should be made of
the effects of that management in order to modify procedures as

required.

 

75

8. In order to conduct studies in national park management,
a staff of university-trained wildlife ecologists should be assembled

and assigned as park biologists.

SUMMARY

Habitat and population interactions of maral (Cervus elaphus

 

moral) were studied in the Mohammad Reza Shah National Park, Iran,
during 1976-1978 in the three vegetation types present: Caspian
deciduous forest, transition zone, and steppe.

Maral densities per 1000 hectares were calculated by the Hahn
method for each vegetation type and season. The maral population thus
was estimated to be 1897 for the park. Transforming the data by the
variate y = log(X + 73.10), significant differences in deer densities
were indicated between seasons, areas, and their interactions.

Using the fecal pellet-group data with a defecation rate of
10 pellet-groups per day, the population estimate was 2096 deer for
the park.

Based on the Hahn figures and pellet-group counts, the average
maral density was 15 and 16.6 maral per 1000 hectares respectively.

The average maral group size was 4.6. The sex ratio among
adults was 27 stags per 100 hinds. Based on summer data, the park
maral had a yearly mortality rate of 13.9% and a life expectancy of
6.7 years. The number of calves produced per 100 hinds was 28.

Single calves were the rule. Gestation approximately lasted about
250 days. The mean date of birth was estimated to be 30 June.
The maral is mainly a grazer, preferring grasses over woody

forages. 0f the 19 plants eaten by the maral, Dactylis glomerata was

76

77

the most-preferred food, the year around. Euonymus sp., and Colutea
persica were highly-preferred winter browse species.

Pellet groups were correlated positively with shrub abundance.
Showing a preference for forest meadow and transition zone habitats,
food consumption was estimated to be at the rate of 2.87 and 3.6 Kg/day
in winter and spring, respectively. The carrying capacity of maral
range was estimated to be 15.6 ha/maral and 11.0 ha/maral for transition
zone and forest meadows, respectively.

More than 8% of the available shrubs were severely hedged in
the transition zone by browsing deer. Hedging and debarking of shrubs
and trees in the transition zone indicated that maral populations there
had reached the range carrying capacity. Maral populations showed signs
of over-abundance. There was an excess of old animals and, in com-
parison with American and European findings, the rate of recruitment
was low. Of all the maral classified, 11.0 percent were in poor
condition.

Wild pigs were important competitors for food. Wild pig and
maral competed heavily for acorns in the fall.

Leopards and wolves were the major predators capable of preying
on maral. Together, it was judged that they killed about 153 or 7.7
percent maral annually. Poaching, however, may have caused more adult
mortality than predators.

Recommendations are made for maintaining maral p0pu1ations in
the national park. It is believed that the excess animals, probably

more than 160, should be removed by controlled shooting.

LITERATURE CITED

78

 

LITERATURE CITED

Andersen, J. 1973. Analysis of a Danish Roe- deer Population
(Capreolus capreolus L.) based on the extermination of the
totalistock. Danish Review of Game Biology, 2: 127- 155.

 

Andersen, J. 1961. Biology and management of roe deer in Denmark.
Terre Vie,. 1:41-53.

Beruldsen, E. T. and A. Morgan. 1932. Notes on botanical analysis
of irrigated pasture. Imp. Bur. Plant Genetics. Herbage
Pub. Ser. Bu11., 14:33-43.

Blaxter, K. L. 1962. The energy metabolism of ruminants.
Hutchinson, London, 329 pp.

Blaxter, K. L., R. N. 8. Kay, G. A. M. Sharman, J. M. .M Cunningham,
and W. J. Hamilton. 1976. Farming the red deer. Department
of Agriculture and Fisheries for Scotland. Edinburgh.

92 pp.

Cadman, W. A. 1967. The management and control of Fallow deer in
the New Forest. Forestry, supplement, 59-63.

Caughley, G. 1967. Calculation of population mortality rate and
life expectancy for thar and Kangaroos from the ratio of
juveniles to adults. New Zealand Journal of Science,
10:578-584.

Caughley, G. '1977. Analysis of vertebrate populations. John Wiley
and Sons, New York. 234 pp.

Chapman, 0. I. 1975. Antlers-—bones of contention. Mammal Review,
3(4):122-172.

Corbet, G. B. and H. N. Southern. 1977. The handbook of British
mammals. Black Well Scientific publications. London.
520 pp.

Cowan, I. 1950. Some vital statistics of big game on over-stocked
mountain range. Trans. N. Am. Wildl. Conf., 15:581-588.

Darling, F. F. 1937. A herd of red deer. London, Oxford University
Press. 215 pp.

79

80

de Nahlik, A. J. 1974. Deer management, improved herds for greater
profit. David and Charles, London. 250 pp.

de Yes, A., C. Eilean, and G. Haghighat. 1977. A master plan for
Mohammad Reza Shah National Park, Iran. Department of the
Environment. Field Document. 46 pp.

de Vos., A., and H. S. Mosby. 1969. Habitat analysis and evaluation,
pp. 135-172 in Wildlife Management Techniques. Edited by
R. H. Giles. ThiFd edition. The Wildlife Society,
Washington, D.C.

Dzieciolowski, R. 1969. The quantity, quality and seasonal variation
of food resources available to red deer in various environ-
mental conditions of forest management. Forest Research
Institute, Warsaw, Poland.

Dzieciolowski, R. 1970. Relations between the age and size of red
deer. Acta therologica, 15:253-268.

Elliott, J. M. 1977. Statistical analysis of samples of benthic
invertebrates. Freshwater Biological Association Scientific
Publication No. 25, second edition. 160 pp.

Firouz, E., E. Etemad, and J. Hassinger. 1971. The wild sheep.
Journal of Hunting and Nature (published in Persian), 135:72.

 

Flook, D. R. 1970. A study of sex differential in the survival of
wapiti. Canad. Wildl. Serv. Rep. Seri. No. 11. 71 pp.

Geist, V., and R. G. Petocz. l977. Bighorn sheep in winter: do rams
maximize reproductive fitness by spatial and habitat segrega-
tion from ewes? Can. J. Zool., 55:1802-1810.

Greer, K. R., and R. E. Howe. 1964. Winter weights of northern
Yellowstone elk, 1961-1962. Trans. N. Am. Wildl. Conf.
29:237-248.

Brieb, J. R. 1958. Wildlife statistics. Colorado Game and Fish
Dept. 96 pp.

Gwynn, J. V. 1965. Deer management: quality versus quantity.
Virginia Wildlife, 26(1):8-9.

Hahn, H. C., Jr. 1949. A method of censusing deer and its applica-
tion in the Edwards Plateau of Texas. Texas Game, Fish and
Oyster Comm. 24 pp.

Harper, J. A., J. H. Harn, W. W. Bentley, and C. F. Yocom. 1967.
The status and ecology of the Roosevelt Elk in California.
Wildlife Monographs No. 16. 49 pp.

 

81

Harris, R. A., and K. R. Duff. 1970. Wild deer in Britain. David
and Charles, Newton Abbot. 112 pp.

Hartley, H. 0., P. G. Homeyer, and E. L. Kozicky. 1955. The use of
log transformations in analyzing fall roadside pheasant
counts. J. Wildl. Mgmt., 19:495-496.

Horn, A. S. 1966. Measurement of overlap in comparative ecological
studies. Amer. Naturalist, 100:419-424.

Jensen, P. V. 1968. Food selection of the Danish red deer (Cervus
ela hus L.) as determined by examination of rumen contents.
DanIsh Review of Game Biology. 5:1-44. ,

Klein, 0. R., and S. T. Olson. 1960. Natural mortality patterns
of deer in Southeast Alaska. J. Wildl. Mgmt., 24:80-88.

Leopold, A. S., T. Riney, R. M. Cain and L. Tevis, Jr. The Jawbone
deer herd. California Fish and Game Dept. Bull. 4. 139 pp.

Lockie, J. D., W. E. S. Mutch, and 8. Cooper. 1967. The Management
of red deer in the hill sheep environment. The Nature
Conservancy. Edinburgh. 68 pp.

Lowe, V. P. W. 1969. Population dynamics of the red deer (Cervus
elaphus L.) on Rhum. J. Anim. Ecol., 38:425-457.

Lowe, V. P. W. 1971. Some effects of a change in estate management
on a deer population, pp. 437-456 in The scientific
management of animal and plant communities for conservation
(edited by E. Duffey and A. S. Watt). Blackwell Scientific
Publications, London.

Mautz, W. W., H. Silver, J. B. Holter, H. H. Hayes, and W. E. Urban
Jr. 1976. Digestibility and related nutritional data for
seven northern deer browse species. J. Wildl. Mgmt., 40(4):
630-638.

Mautz, W. W. 1978. Sledding on a bushy hillside: The fat cycle
in deer. Wildlife Society Bulletin, 6(2):88-90.

Mech, L. D. 1970. The wolf: The ecology and behavior of an
endangered species. The Natural History, New York. 384 pp.

Mitchell, 8., D. McCowan, and I. A. Nicholson. 1976. Annual cycles
of body weight and condition in Scottish red deer (Cervus
elaphus). J. 2001. Land. 180:107-127.

82

Moran, R. J. 1973. The rocky mountain elk in Michigan. Research
and development report. No. 267. Michigan Department of
Natural Resources. 93 pp.

Mueller-Dombois, D., and H. Ellenberg. 1974. Aims and methods of
vegetation ecology. John Wiley and Sons, New York. 547 pp.

Neff, D. J. 1968. The pellet-group count technique for big game
trend, census, and distribution: A review. J. Wildl. Mgmt.,
32(3):597-614.

Peek, J. M., A. L. Lovaas, and R. A. Rouse. 1967. Population
Changes within the Gallatin elk herd, 1932-1965. J. Wildl.
Mgmt., 31:304-315.

Petrides, G. A. 1975. Principal foods versus preferred foods and
their relations to stocking rate and range conditions.
Biol. Conserv., 7:161-169.

Petrides, G. A., and U. de V. Pienaar. 1966. Availability and
selection of prey by six large predatory mammals. Mimeo-
graphed Report, Michigan Polar-Equator Club, U.S.A. 2 pp.

Pimlott, D. H., J. A. Shannon, and G. B. Kolenosky. 1967. The
interrelationships of wolves and deer in Algonquin Park.
Trans. Northeast. Wildl. Conf. 16 pp. Memo.

Riney, T. 1957. The use of feces counts in studies of several free-
ranging mammals in New Zealand. New Zealand J. Sci. Technol.
B38(6):507-532.

Riney, T., J. S. Watson, C. Bassett, E. G. Turbott, and W. E. Howard.
1959. Lake monk expedition. New Zealand Dept. Scient. and
Indust. Res. Bul. No. 135. 75 pp.

Robbins, C. T., and A. N. Moen. 1975. Composition and digestibility
of several deciduous browses in the northeast. J. Wildl.
Mgmt., 39(2):337-341.

Robinette, W. L., J. S. Gashwiler, J. B. Low, and D. A. Jones. 1957.
Differential mortality by sex and age among mule deer.
J. Wildl. Mgmt., 21:1-16.

Schaller, G. B. 1972. Serengeti Lion. University of Chicago Press.
Chicago.

Schaller, G. B. 1977. Mountain monarchs, wild sheep and goats of
the Himalaya. University of Chicago Press, Chicago. 425 pp.

83

Shafer, E. L. 1963. The twig-count method for measuring hardwood
deer browse. J. Wild1.Mgmt., 27:428-437.

Staines, B. W. 1970. The management and dispersion of a red deer
population in Glen Dye, Kincardineshire. Ph.D. thesis.
University of Aberdeen.

Staines, B. W. 1974. A review of factors affecting deer dispersion
and their relevance to management. Mammal Review, 4(3):79-91.

Taber, R. D. and R. F. Dasmann. 1954. A sex difference in mortality
in young Columbian black-tailed deer. J. Wildl. Mgmt.,
18:309-314.

Taber, R. D. and R. F. Dasmann. 1958. The black-tailed deer of the
chaparral. Calif. Dept. Fish and Game. Bulletin No. 8.
163 pp.

Telfer, E. S. and G. W. Scotter. 1975. Potential for game ranching
in boreal aspen forest of western Canada. J. Range Management,
28(3):172-l80.

Ullrey, D. E., W. G. Youatt, H. E. Johnson, L. D. Fay, and B. E. Brent.
1967. Digestibility of cedar and jack pine browse for the
white-tailed deer. J. Wildl. Mgmt., 31(3):448-454.

Ullrey, D. E., W. G. Youatt, H. E. Johnson, P. K. Ku, and L. D. Fay.
1964. Digestibility of cedar and aspen browse for the white-
tailed deer. J. Wildl. Mgmt., 28(4):791-797.

Van Den Brink, F. H. 1968. A field guide to the mammals of Britain
and Europe. Houghton Mifflin Company, Boston. 221 pp.

Verseshchagin, N. K. 1967. The mammals of the Caucasus. A history
of the evolution of the fauna. Translated from Russian by
A. Lerman and B. Rabinovich. Israel Program for Scientific
Translations, Jerusalem. 816 pp.

Whitehead, G. K. 1950. Deer and their management. County Life
Limited, London. 370 pp.

Whitehead, G. K. 1972. Deer of the world. Constable, London.
194 pp.

Zuckerman, S. 1953. The breeding seasons of mammals in captivity.
Proceedings zoological society of London, 122:827-950.