The Asiqn elephonf (Efeph as mqximus): its hqbitqt, stqlus qnd distribution in

Arunqchql Prodesh, Indiq

Bharath Sundaram . Surendra Varma, Arun Venkataramanand Raman Sukumar

Abstract The sun'e)'

and status

of

assessed forest

elephants in some areas

contiguitl', distribution

of Arunachal

Pradesh

Tirap and Papam-Pare had high, medium and low abundance

areas respective\r Qualitative information on district wise

to identifl' likely gaps in the protected area network. The

elephant status, distribution and conservation is also

survel, also examined changes seen in elephant distribution

between 1984 and the present. The study was carried out
through field and questionnaire surve)rs in the administrative
districts of the state. Elephant habitat maps with vegetation
types and elephant distribution information were produced.
The encounter rates of elephant signs per kilometre were
calculated to ascertain specific zones of high, medium and
low elephant abundance. Based on the results, East Kameng,

presented. The survel' found very patchy habitat in many

areas and many areas that are important for elephants do
not fall under the protected area network. The paper also
discusses specific recommendations for conserving the
species and its habitat.

Keywords Elephant, Assam, Arunachal Pradesh, Humanelephant conflict.

fntroduction
Arunachal Pradesh, the largest state in the north-east India,
covers an area

of

83,743 km2 between 26' 28'-29" 30'

and

97" 30'-97" 30' E. Lying in the Eastern Himalayan region of

the Himalayan biogeographic zone, Arunachal Pradesh is
located at the junction

of

the Palaearctic and Oriental (Indo-

Malayan) realms (Rodgers

&

Panwar, 1988). Arunachal

Pradesh comprises mountains that ring eastern Assam, terrain

thrown up by the sharp twisting of the Himalayan ranges as

they turn suddenly from a southeastern to a southerly

direction. These mountains drop precipitously into the
tropical lowland evergreen forests of the region. Because

of

the wide gradients in elevation (50 m-7,000 m asl) and

annual rainfall

(1

,000 mm-5,750 mm), an a:ray

of forest types,

ranging from tropical evergreen forests, tropical semievergfeen forests, riverine semi-evergreen fotests, subtropical pine forests, temperate broadleaved forests,
temperate coniferous forests, alpine forests, and high
montane grasslands are seen. Such a diversity of habitats
harbouring rich plant and animal life, packed into a relatively
small are4, makes Arunachal Pradesh a hotspot of global
biodiversity. It is also home to over 5,000 species of plants
(including 600 species of orchids), 500 species of bitds and
29 species of mammals listed under Schedule I of the Indian
Wildlife Protection Act. These include 4 species of latge
cats, 7 species

of

primates, and 3 species

oI

goat antelopes.

There are over 100 species of amphibians and rePtiles

recorded from the region (Anon. 1994).
Arunachal Pradesh shares international borders with Bhutan
in the west (border length 160 km), China to the north and
northeast (1,080 km), and Myanmar to the east (440 km).
Culturally diverse, Atunachal Pradesh, with a population
under 10lakh (of which 70% are tribal people), has 21 mafor
tribal groups with over 100 e thnically distinct subgroups and
over 50 distinct languages and dialects. Spread ovet 14
districts, the people of Arunachal Pradesh practice seminomadic swidden agriculture (also cilIed jboom cultivatioo),
terraced wet agriculture, high montane pastoralism, and
traditional trade and barter. Arunachal Pradesh is a Iand of
mighq' rivers such as Siang, Dibang, Lohit, and Kameng, all
of which ioin the Brahmaputra in the Assam valley Over
70'h of Arunachal Pradesh is forested, md 17-8oh of the

total land area falls under the protected area network (Fig.
1), with one Biosphere Reserve (BR) (Dibang-Dihang
Biosphere reserve), one Tiger Reserue (fR) Q"lamdapha Tiger
Reserve), one National Park (IrlP) (Mouling National Park),
10 Wildlife Sanctuaries (WLS) and one Orchid Sanctuary
(Sessa Otchid Sanctuarl). Eighty-six Reserved Forests (RF's)
account for an additional 72.7'/o o[ the total area. The rest
of the forested areas come under the pun'iew of the 3,649
villages in Arunachal Pradesh and are classihed as Village
Forest Resen'es (VFRt) or Unclassed State Forests (USF's)'

Bharath Sundaram, Surendra Varma,

ArunVenkatatarr:.aln & Raman Sukumar

At least tw'o populations of the Asian elephant (Ehphas

nainns) extend along the Himalayan foothills and plains

Asian Elephant Research and Conserr.ation Centte

(A division of the Asian Nature Conservation

from northern \n'est Bengal eastwards through Assam, Bhutan

and Arunachal Pradesh (Sukumar, 1989). It has also been
estimated that 10,000 km2 of hill1' habitat may be available
for elephants in Arunachal Pradesh alone. The Arunachal
Pradesh trorest Depattment estimates the population of
elephants in the stete at 4,000, u'hich is twice that o[ an
earlier estimate (Lahiti-Choudhurt', 1980). This discrepancv

Foundation)

Centre for Ecological Sciences (CES)

Indian Institute of Science (llSc)
Bangalore 560

(11'2

India
Galah 222

flulv

2003)

43

44

thc lact that elephants movins into r\rlrnachal

and s'hen thel occurred. Proximit_r' to the closest human

from z\ssam u'cre also counted (AESG Rcport, 1981). In

c<-rllaboration u'itlr the Forest Department, the Task Irorce
of the IUCN/,\sian Elephant Specialist Grcrup (Lahiri-

settleme nts u'as also noted. [n addition to ftelcl data collection.

Choudhurl', 1980) mapped the distribution of elephants in

northeast India (Lahiri-Choudury et a/., 7984). Hou,er.er,

included information on distribution, movement patterns,

and numbets of elephants. Since verl' ferv elephants in
Arunachal Pradesh are resident, data on seasonalitv of
sightings/mo\rement rvere also collected. Fieldrvork was
carried out during the dry periods of 2001.

ct

'ulcl

l>c clue k r

rvidespread destruction of foothill forests through

developmental and other anthropogenic actir.ities have
shrunk the range of the elephant, consequently restr'ct ng 't

to smaller landscapes having very little connectivit)'. An

increase of encroachment in forested areas has also led to
an increase in elephant-human conflict.
The prime objectives of this survey were to assess the forest
contiguit)' and status of elephants, to identify possible gaps
in the protected area network in some areas of Arunachal
Pradesh, to examine the changes seen in elephant distribution
betweeo the 1984 stud)' and the present one, and ta proiect
all the data on maps that could be used by both the Forest
Department and researchers for better managemenr and

further research.
Methods

The status and habitat available for elephants were

determined through field sun'eys and questionnaire sun'e)'s.

Field survelrs were undertaken in East Kameng,

$57est

I(ameng, Papum-Pare, Lower Subansiri, Dibang Vallel', Lohit,

Tirap and Changlang districts of Arunachal Pradesh. Survey
of India toposheets were used to assess forest contiguitf in
the low- and mid- elevation (100-900m) areas. Information

regarding the area was first gathered from the Divisional

Forest Office, after urhich animal trails in the forest were
identified. These trails were then traversed on foot, and data

on elephant signs, i.e. dung, pad marks, scratch marks, and

feeding signs were collected. GPS locations were taken at
kilometre intervals, and the habitat at that point was classified
by noting geographical features, and the fout species of rees
characteristic of the vegetation in the nearest vicinity were
noted. Direct sightings of elephants were also recorded as

Table 1 List of

areas where

seconclarf infcrrmation rvas gathered through questionnaire

surre ):s in villages and Divisional Forest Offices. This

Encounter rates (freqyency of sighting elephant signs per

kilometre traveled on foot) were calculated ffable 1). From
these encounter rates it was possible to determine, within
the study area, specific zones of high (encounter rate in the
range 4 and above), medium (encounter rate in the range 2.5
to 4), and low (encounter rate in the range 1 to 2.5) elephant
abundance.

Maps wete created using Maplnfo Professional, ArcView

GIS, and GIWView. La1,s1r .1 political boundaries, prorecred
atea network, forest cover were created using VMap Level 0
data (the data set was derived from 1-km resolution Advanced
Verl' fli*1l Resolution Radiometer (AVHRR) data spanning
a 12 month period, from April 1992 to N{arch 1993), and
Global Land Cover Chartcterization datasets obtained from
the International Steering Committee for Global Nfapping,
Geographical Sun'e)' Institute, Japan. The GPS sur\re)' points
'\vere overlaid on the forest co\rer map (Fig. 2), and infetred

elephant distribution (from secondarl, information) was

plotted (Fig. 3).
Results
The elephant ranges in Arunachal are mainly distributed along
the foothills adjacent to Assam. Elephants are found usually
between the altitudinal range from 2,000 to 6,000 m asl but
there is some evidence to show that they could migrate to
higher levels in seatch of food and other resoutces especially
during v/inter months. A total of 1,600 elephants were
estimatcd for the state during 2001 elephant census (Forest

ground surveys for dung were carried out, with encounter rates for each area

Distance Number of
walked ftm) signs

Encounter

L.rr.l

Bandardewa

RF, USF-

l2

2.33

Chessa Papum-Pare Bandardeu,a

Kimin
Papum-Pare Bandardewa
Deomali Tirap
Deomali
Deomali Tirap
Deomali
D'Ering East Siang Pasighat

RF, USF
RF, USF.RF, VFR
RF, VFR
WLS

I]

1.88

15

24

1.6

Khellong
Khellong

Area

District

Chessa Papum-Pare

lP'

Jellosa

llas t

Kameng
East

I\ameng

Iorest ljlvlslon

Protection

28

nte/km

15

79

5.21

29

4.14

l6

88

5.5

WLS

12

78

6.5

\(1LS

15

89

5.93

()ajah 2z: ULrrv z(ru:)

45

Departmer.rt, unpublished teport). \\'ithin Arutracl'ral Pradesh,

elephants occur in Papum-pare, Lorver Subansiri, East Siang,

r\ssam <lue
cultivation.

to the

large-scale trlnsformation oF land fnr

East &West Kameng, Lohit Dibang Vallev, Tirap &

l>apant-Pare and I -oaer

Changlang districts.

East and lVest Kameng disticts

Areas surveted in these nvo districts incluclc Pakhui \\'t,S

and Iihellong Forest Dir-ision, both

of

rvhich support

abunclant elephants and experience severe hun-ran-elephant

conflict. The high abundance mal be due to these areas
bordering Nameri NP and Sonitpur RF in Assam, from where
elephants tend to mo\re out. \\'hile the forest t1'pes in Pakhui
\\ILS range from semi-evergreen to evergreen *'ith Patches
of secondarl'jbooru forests, the forests in the Bhalukpong

side (I(hellong Forest Division- Amartala, Doimara and

Papum RF's) are mostl)' degraded, u,ith some patches of
evergreen forest still existing due to their inaccessibilitrr
Elephants range right from the foothills of the Dhansiri rive r
(at the Bhutan botder), to Papum RF east of Pakhui V'LS. A
specific arez of high human-elephant conflict is Tipi (a town
close to Bhalukpong). The position of the Tipi Orchid
Reseatch Centre (situated

increased levels

in a natural elephant

Path$'ay),

of human encroachment, and

the
construction of a Territorial Range Office on an elephant
path have been the reasons for increasing levels of human
elephant conflict. Project Elephant has recognised the
I(anreng -Sonitpur interstate luea (ca 4,300 km'z) as an
Elephant reserve Q'ilational Elephant Consetvation Action
plan, 1992), but this area experiences a lot of pressure in

,fabansii Di.rtids

The areas survel'ed in these t'wo districts include the Itanagar

\flLS and Bandatderva Forest Division, which had medium
highly migrant animals' These areas
elephant abundance,
"vith
are also highly ftaumented and degraded due to human
encroachment and illesal deforestation. r\reas such as Poma,
Jote, I{imin, FIoi, Tataiuli, Chessa, Hollongi, Changmara,
I{okila, Tengabari, and Baliian have a matrix of habitats
composed of semi-er-ergreen ftrrest, evergreen [<rrest, and
.cultivation. Unplanned development in the Itanagar area has
also destrol'ed much of the habitat and has been identified
as an issue of concern in 1984 (Lahiri-Choudhurl" 1985).
An elephant reserve has been planned in Papum-Pare district,

but the success of this plan depends entirely on how

effectively' encroachers can be removed' This area shows
vety little influx of elephants from Assam because the areas
on the Assam side ate completely under cultivation.

In the Subansiri

range

of Panir RF (Bandardewa Forest

Division) disturbance affecting elephants has been observed.

Dolunmukh is a village botdering Panir RF, which has not
reported the presence of elephants for the Past 10 years due
to such disturbance. Though the disturbance has ceased a
few 1'ears ago, elephants are )'et to return. These areas have
large patches of semi-evergreen and evetgreeo forests. Mote
interesting\., Dolunmukh was reported to have been an area

of high human-elephant conflict

1 Eagles Nest WLS

2 Sessa Orchid Sanctuary

3 PakhuiWLS
4 ltanagar WLS
5 Kane WLS
6 Yordi Rabe Supse WLS
7 Mouling NP

I
I

Dihang-Dibang BR
D' Ering Memorial WLS
'10 Mehao WLS
'11 Kamlang WLS
12 Namdapha TR
Wls- Wildlile Sanctuary
NP- Nalional Park
BB- Biosphere R6erve
TR- Tiger Heserve

G{rh

22: (fulr' 2003)

(Lahiri-Choudhuty et al.,

46

1984). A maior hldroelectric project

ofa dam across the Subansiri

I-nhit

involvins

rhe

consrrucrion

Dirtitt

The floodplain areas of the Lohit River, such as the Paya

and Digaru Ranges, were surve)'ed in the Lohit Forest
Division within this disttict. Of the 7 ranges here 2 showed
seasonal presence of elephants. Dung encounter rates
indicate that these areas have medium elephant abundance.

Affected heavily by flood, the landscape in these areas is

highly amorphous. Struck by a major earthquake in 1950,
many

of the natural forests of this area were destroyed due

to inundation and the changing of the course

of

Saccharam grasslands and plantations.

Plantations were established following the earthquake to

restote some of the original fore st. In providing fresh herbage
for elephants after the first rains, this area assumes immediate
importance and hence it is necessary to bring these areas
undet the protected area network. This area is also heavily

disturbed due to the presence of 'kbatis' (herds of

unproductive catde). Overgnzingwas seen the Digaru area.
The Paya and Digaru areas are approximatell' 300 km2 in
size, and this area supposed\'has at least 70-!2 klturts @a,

Di.;tict

The areas in and around Nfehao WLS were surveyed. N{ehao

WLS (281.5 km'Z) lies along an elevational gradient and
evergreen forests are found up to 900 m (an area o[ around
100 km']). This area shows medium elephant abundance, and
appears to be highly disturbed with a high degree of
encroachment (especiallf in the I(oronu and lppipaani areas),
where the people are not aware that they are living in the
midst of a wildlife sanctuarlr. Elephants that use the DibruDeomali elephant corridor sometimes visit this. area.
Changlang

Dirtict

Namdapha TR (1985 km) is located on the Myanmar border,

and movement of elephants to and from Myanmar cannot
be totalll' ruled out. This area was found to have low elephant
abundance. A survey by our team in 1999 also yielded very
poor results as far as elephants were concerned. The forest

types

in Namdapha TR range from

semi-evergreen to

evergreen on the lower reaches. Evergreen forest habitat has

been found to be sub-optima for elephants and this may be
the reason for their low abundance.
T;raP Dirtrict

of

catde). Such large number of cattle is

bound to put undue pressure on the grassland ecosystem,
2500-3000 heads

11/e1,

the Lohit

River. The habitat in this area is a combination of secondary

riverine forest,

Dihang I

is also being planned.

and should be brought under control. Choudhury (1999)

has also cited illegal capture of elephants to be common in
this district, but no such instances u/ere observed during this
study.

Fiq.

lClosed Canopy Foresl

E Open Canopy Forest
E Agdculture

Extent

of

In this district, the Deomali Forest Division (286 km2)

was

surveyed. The Arunachal Pradesh Forest Corporation used

this area for timber operations until 1997 and selectively felled
patches are recuperating. This is an area of high elephant
abundance, with elephant influx from Nagaland in the west, as

forest cover in Arunachal Pradesh

50

50

Krlometers

()aiilt 22: (fulv

2003)

47

well as movement of elephants from neighbouring Joypur

RF in Assam. A large number of natural salt ltcks (pungs)
were found in the Namsangmukh area. These areas are
classiFred as VFR's and RF's, and must be incorporated into
the protected area network, taking into account their value

Discussion

for elephants. Elephants also frequently move from Tinsukhia

and Digboi areas (especially around Dibru-Saikhowa NP) to
this area and vice-versa

habitat. Mosaics

East SiangDis*ict
The D'Ering Memorial \flLS was surveyed in this district. The
D'Ering \fLS (184 km2) is composed mainly of Vettiaeia
grasslands and plantatiorc of Dalbergia znd Bombax. It is
of high elephant abundance and is used
another
^teeseasonally by elephants (before and duting the monsoons)'
The zrea is also flood prone and is rendered a virtual island
during heavy rains and flood. Heavy siltation occurs in the
areas affected by flood. Elephants frequendy ctoss over from
Jonai and Kobo Chapori in Assam to this area. Cited as an
important movement path for elephants (Choudhury, 1999),
the Dibru-Deomali elephant corridor includes D' Ering WLS,
Jonai and Kobo Chapori areas (Assam), Dibru-Saikhowa Nl
Joypur RF and Deomali Forest Division. Consequendy, this

whole area (ca 4000 km2) is under the Dibru-Deomali

Elephant Reserve. Presendy D' Ering WLS is not protected
effectively due to the shortage of manpower and resources.
As a result many people from Pasighat and Mebo use it as a
hunting gtound. Frequent burning of the grassland also occurs.

Habitat patchiness and Elepbant-Haman Conflict

Most of the areas surveyed, excepting

of forest

a few, showed very

patchy

types such as semi-evergreen'

degraded thoomed forests, degraded gtasslands and degtaded

riverine forest are interspetsed uzith cultivated lands. During
their attempts at maintaining normal ranging pattem, elephants
encounter an increased interface between natural vegetation
types and cultivation. Elephant-human conflictwas a common
problem in most of the areas surveyed. The people of

Arunachal Pradesh have always been livingin close proximity

to their forests, but uncontrolled development has already
destroyed many prime elephant ateas, such as the foothill forests
in Papum-Pare district and Lower Subansiri. The problem of
elephant-human conflict is compounded by the fact thattlarge
proportion of agdculture in Arunachal Pradesh is subsistence
agriculnre. Crop depredation affecs the subsistence agdculturist
very hard, and the existing system is not equiPPed to deal with

this problem. The formulation and implementation of a

compensation scheme will greatly help improve relations
between the common people and the governmental agencies
involved.
Gaps in

tbe

pmtected arca rctu'ork

Many areas that are imPortant for elePhants do not fdl under
the protected area network. The Deomali area in Tirap district
is one such example. An important link in the Dibru-Deomali
corridoi, this area is Presendy classified as a VFR. It is not only

important to bring some areas under the Protected area

network, but existing Parks and sanctuaries must be well

protected as well. The D' Ering Memorial \(ildlife Sanctuary is

for elephants as they use the grasslands
a very imPort^At
^tea
extensively for feeding. Limitations of manporver and resources
make the job of protection difficult, and a solution to the

problem lies

Fig.

in finding a cost-effective temedy to ensure

Distribution of Asian elephants in Arunachal Pradesh based on a questionnaire survey

'l HrghElephanlAbundance
EElephant Orstribution
aSurvey Points

Gajah 22: (fulv

2 Medrum Elephant Abundane

3 LwEleohantAbundance

50

50

Kilometers

48

protection bv

inr'<--,h.'ing

thc local communities. Since the

eleplrant populations in the northeastcrn region knorv no

political bounclaries, it is necessan'to ensure their protection in
the neighbourin.q states of Assam, Nleghalaya and Nagaland.
Unprecedented levels oIdeforestation in Assam often cause
problems in the areas where cross-border moven-rent of

elephants is common, as for example

in the Kameng

@orderingv-ith the Sonitpur and Balipara forests of Assam)

and East Siang-Tirap @ordering with Dibrugarh-Dieboi areas)
zones. The Forest Department in these states must \Mork out
a common strategl' f61 1le protection of elephants on a regionwise scale b)'identitpng these ke)' areas.

Fig.4 The location o[ K

cbsd

orn

rertunurc

ap;"

Prutnntion ol kqy ltabitats

Habitats such as the riverine semi-evergreen forests and

grasslands ate extremelv important for elephants, as thel,are a
readl' source of forage, e special\' following the first rains. These
habitats are constantl), under thteat due to possible conversion

to cultivated land. A viable alternative to livestock grazing is

stall-feeding and this must be promoted in a large scale in
order to reduce the pressure on forest resources. It is very clear
that there can be no long-term solution u'ithout the combined
elfort of the Forest Department and the indigenous people
of the area.

ng Sonitpur area tn the Arunachal Pradesh and Assam

canopy Focst

canopy Facst

Fig. 5 Map of the Dibru-Deomali zrer of Arunachal Pradesh and Assam

D DDrusdkhda NP
a cbsed carcpy Fde{
E open canoDt Foresl
O &nulue

I eopun toerent

Gaiah Z2: (luh' 3)t).D

49

Acknowlcdgements

Refercnces

Thc surve). rvas made possible through the support provicled

by the forest officials in Arunachal Pradesh and t\ssan-r. We
would like to extend our thanks to Nlr. S. S. Choudhurl',
Chief Conservator oI Forests (Wildlife), N{r. S. Deoti,

Asian Elepbant Specialist Group. 1,987. Pronedings of tbe First

nt e e ti ng of N o rt b -E a s t 7'a s k fi rn, A.ri a n E leP h a n t .fpe ci a li r t G ro ap,

oI Forests (Arunachal Circle), Deputl'

of Forests, Mr. D.N. Singh, N{r. C, D. Singh,

Anonvmous. 1994. The Wildlife (Protection) Lct 1972. (2"r

Edition). Natraj Publishers, Dcl.rra Dun, India.

A.I(. Sen, Mr. C. Lorna, N{r. Umakant, TatunJohri, X{t.

Santhosh Kumar, Mr. Sitang, Mr. H.B. Abo, IVr. N{odi, N{r.
D.Shra, N{r. Gupta-Choudhary (Assistant Field Director) and
N{r. S.S. Chandiramani (X.esearch Officer) for their help during
the project.

Choudhuryi A. 1999. Status and consen'ation of the Asian

Elephant E/ephas ntaimu in north- eastern lndia. Mammal
tufiew 29: 141-173.

Conservator
Conservators

IUCN/SSC.64 pp.

N{r.

Lahiri-Choudhury, D. K. 1980. An inteim report on

We like to thank Dr. K. Harikaran, Systematic Botanist, State
Forest Research Institute, Chimpu, Prof. P.C. Bhattacharjee,

and distribaion

of

elepbants in nortbeast

tbe statas

India. IUCN-SSC Report.

16 pp.

Guwahati Universitl', Mr. Richand Ahmed, Extra Assistant

Honorary Wildlife Warden, Mr. Nabm Bate, Membeg State

Lahiri-Choudhury, D. K., Dey, B., Sar, C., & Mukheriee, A.

1984. Srnnaised report on Elepbant Habitat Sanry in Aranacbal

tVildlife Advisory board, Mr. Vivek Menon, Executive

Pradesb.

Commissioner, Murkong Selek, Mr. Jose B. Nabum,

Directot, Mr. Sunil Subba, Field Officer, Northest Region,

Wildlife Trust of India. Ms. Tanushree Biswas. Ms.
Padmawathe Ramakrishnan. Wildlife Institute Of India and
Mr. N. Narendran, Mrs. Nirmala and Mr Palani of Centre
for Ecological, Sciences, IISc, for their inputs and support
at various stages of the surveyr \l(e would also like to thank
Dr. T. V. Ramachandra, CES, IISc, Mr. N. Samba I(umar
WCS (India program) for GIS support and georeferencing.
The survey was jointly executed by Asian Elephant and
Research and Conservation Centre, Bangalore and Wildlife
of India, New Delhi. 'Jfle acknowledge the sen'ices

Trust

rendered by the staff members of both organizations.

IUCN/WWF Report. 21

pp.

Lahiri-Choudhury, D. K. 1985. Distribation, popttlation estimah,

and .rtatas of elepbant in North-East India. IUCN-SSC Report.
1o pp.

N{inistry of Environment and Forests, Government of India.

7992. National Ehphant Conrcruation Action Plan- Exemtiae
Sanmarl. 22 pp.
Rodgers,

!(A. & Panwar, H.S. 1988. Planninga wildlfeprotected

area nehtork in

India.2 vols. Project FO: IND/82/003.FAO,

Dehra Dun.
Sukumar, R. 1989. The Asian Elephant: Ecology and Management.
Cambridge University Press, Cambridge.

A bull in musth. (photo: c()urtes\.Dr. tl.I.b,. liatugaha)

Gtjth 22: (fuh'

2t){13)