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BIOLOGY OF SAMBAR DEER (Cervus unicolor Kerr, 1792) IN NEW ZEALAND WITH PARTICULAR REFERENCE
TO DIET IN A MANAWATU FLAX SWAMP
A thesis presented in partial
fulfilment of the requirement for the degree of Master of Science in Zoology at
Massey University
SIMON DOUGLAS KELTON
SAMBAR HIND, MOUTOA (L. Barnard)
i i
ii.i
ABSTDACT
This MSc. thesis is the first detailed account of the biology of sambar deer (~:~-~ l!..l!Ic;.£~;;,2£ Ker:c, 179~n in New Zcalando Observations ware made for fifteen months on a.pproximo.tcly 35 animals inhabiting flax swamp at Moutoa, southern Manawatu.
Sambar are shy and cautious, mostly nocturnal and prefer dense cover. Methods used to overcome problems of
direct observation of sambar are described and their relative effectivenesses are compared. Habitat requirements are also discussed.
Diet of Moutoa deer was determined by ana1ysing monthly faecal samples supported by direct observation of feeding and examination of feeding ev:i.dence., A quantitative me·thod of faecal analysis based on the a:rea of pla.nt cu.t.icle
present. is described .. Seasonal changes in diet composition are detailed. Flax and rank grasses comprise the greatest proportion of the diet, wh).le rye grass was present in low proportions and clover was absent from the faeces. Evidence is presented suggesting sambar dee:c do not compete-! \·.ii.th domestic stock for high quality forage.
The eight most frequent forage species f ou:nd in faeces were sampled seasonally and analysed for Acid Deter- gent Fibre, energy, nitrogen (crude protein) and water
content. There was no correlation between changes in forage quality throughout the year and seasonal changes in dii~t
composition. Other factors involved in forage selection are discussed.
The Moutoa breeding population is viable and had an estimated average age structure of 3~ adult males,
4&%
adult females and 18% juveniles.
In New Zealand sambar appear to breed throughout the year with two peaks of tncreased rutting ac'c,lvity in
J~ne, July and August, and in November.
Evidence is presented that the majority of stags shed their antlers annually, in contrast to the previ.ously
·accepted belief that they hold antlers. for two or more
years. Antler-cycle is closely associated with the breeding
iv
cycle, most sightings of stags in hard antler occurring from J·une to Novembero Examination of available information on breeding and antler cycles in Australian sambar revealed similar cycles to these in New Zealand, whereas in India i t appears a single peal-;: in rutting occurs from October to December, with a corresponding antler cycle.
Herds are loosely struct~red and generally comprise small family groups t commonly a hind, yearling and favmo
Young stags generally form groups of two to four individuals while old stags evidently lead solitary lives except in the breeding season when they were often observed with one or two hinds,, Rutting stags a.re territorial with olfactory and visual cues apparently serving to exclude rivals •. Hoarlng or fightin.g, apparently common in India. during the rut, is rare in New Zealand.
Evidence is presented that some hybridisation with rusa dear (~~ ~ Blainvillef 1S22) occurs in
the Bay of Plenty. A comparison of cranial characters between Manawatu and Bay of Plenty deer was inconclusive because of :i.nsufficient· numbers of slmlls. Sexually dimorphic cranial characters are given for animals from the Manawatu area.
Sarubar deer skulls were aged by counting the number
or ·
annuli in the cementum pad of molarif orm teetho
ACKNOWLEDGEMENTS
My supervisor P Dr J e Pe Skip\':orth, deserves my sincerest thanks for his constructive· help during all of
my research. and the3is prepare.tion.
Sincere:: thanks are also due to Mr and Mrs A .E.
Hunter, who managG Mo"J.toa. Without their enthusiastic help and kind hospitality this research would not have been possible.
The Hew Zealand Forest Service provided $500.00 towards my travel and often extended their generous help and advice for which I am grateful. In particular I apprec- iate the help and information given by Dr Co Challies,
v·
C.L. Batchcler and M. Douglas (Forest Research Institute, Islam), I. Logan, W. Simmons and J. Sut.ton (Forest Service, Palmerston North), and H. Vipond, ,Judy Gamble and J. Knowlton
(Forest Service, Rotorua)e
Considerable help was also given me by Me Daniel and Dr M. Rudge, (Ecology Division, D.S.I.R.) who gave me the benofit of their own fleld experience.
The following people also made their own unpublished data available to me : M. Daniel on Rot.orua sambar ; Dr Mo Draisma, (Deer Advisory Council of Victoria, Australia) on breeding and antler condition of Australian sambar ; J. Rudd
(Deerstalkers' Association, Rotorua) on breeding and antler condition of Rotorua sambar.
I am indebted to Dr G. Arnold, D. Drummond,
R. McCammon, P. McGr·cgor, Dr E. Minot, Dr T. Moore (all of Massey University) antl C.L. Batcheler for statistical
assistance and advice on craniometric analysis. In addition Dr E. Minot.kindly ran the "Genstat" program for analysing my nutrient data.
Thanks are also due to the following Massey staff : Dr A., Davey for allowing me the use of nutrient analysis equipment, and Mrs Y. Moore and J. Raven for showing me the use of this equipment ; Dr ·E., Kirk who provided a diar.ion.d saw for tooth sectioning ; Jo Cloustcn and staff of the Central Photographic Unit for prepar~tion of photographs ;
'Vi
and the illustrator, K. Korn0.9rffor~ for the use of the Kroytype lettering machine.
I . appreciate the invaluuble informatj.on concerning
practical experience with sambar given by C. Fergusson, J. Hargraves, A. Harnett, O. Maher, L. Rowe, R. RoNep · Le Shailer, M. Shailer, P. Smith and Wo Wallace.
I also appreciate the help given me by Dr R.A.
Fordham and Ie McKelvey for -the preparation of the skeleton shown in Plate 15 ; I. 'McKelvey for helping me erect J~he
Massey hide ; L. Barnard for the use of his photographs shown in the frnntispiecc, Plate 4 and Plate 16 , and f 01·
helping erect and supplying material fol' the Coffin hide. My thanks to N. Haack, P. Lo, Adrienne Pease, Mr and Mrs D.C. Pease, Moyra Roberts and my parents who assisted me in many ways and provided encouragement on innumerable occassions.
Some additional help with the preparation of the script was provided by Dr I. Stringer and Professor B.P.
Springett.
Adrienne Pease and my parents helped by proof reading.
Lastly many thanks are due to Y.rs J. Evans and Mrs F .S.e Wicherts tor the setting out and typing of this thesis.
TABLE O~ CONTENTS
ABSTRACT
ACKNOWLEDGEMEi·lTS
TABLE 01<" CONTENTS
LIST OF TABLES LIST OF FIGURES LIST OF PLATES
PAR'£ I CHAPTER 1
1 .. 1 1.2
1.3
lo~'-
1.5 1.6
PART II
CHAPTER 2 2.1
2.2
2.3
CHAPTER
3
J.13.2
INTRODUCTION
Introductory Description 'l'axonomy
Distribution Mout.oa
The Flax Reserve (Main Study Area) Climate
DIET
OBSERVATION
Direct Observation
2.1.1 Introduction to methods 2.lo2 Da.ytime observations 2.1.3 Night observations
Indirect Observation (Feeding Sign) 2. 2.1 Flax
2.2o2 Poplar and willow
2.2.3
Grasses Feeding HabitsFAECAL ANALYSIS
Introduction Methods
3~2.1 Collaction of faeces J.2.2 Initial treatment
v vii
xi xii xiv 1 1 1
3 5
11
13
1$
19 19 19 19 21 21 24 24 25 2$
2$
32 32 35 35 36
J.2~3 Nitric 8cid method of cuticle
preparation 36
CHAPTER
4
PART III
CHi~PTER 5
5.4
,3.2.I+
3.2 .. 5 3.2.6 3.207
Hesults 3 .3 .. 1
3.3.2
Reference cuticle colJection Recoznition ite~s
Analysis procedure Hurr1en sariples
Rurr:en analysis Faecal analysis
NUTH.TENT ANALYSIS
Introduction Forage QUC:ility
4 ,, ., • -=.. • ...!.. Defining forage
4.2.2
Protein11-.2.3 Energy h. ~~ ~ lt Fibre
4.205
r:oi~-t'...<re content Sampling TechniqueAnalytic al !'-~et hods
qu~li·'· c;. - (., v J
l~. 4 0 1
4.Lt.2
! 4 "
. -+. • )
Crude prot8in deter~ination
Gross energy determination Acid Deter~ent Fibre (A.D.F.)
det e~inat ion
?-:oi st ure cont er.t det erir.inat ion Results
POPULATION DYNiJ-:ICS
Populat io1~ Size
Sex and Age Composition Breeding Biology
5.3.1 India and Sri Lanka
5~3·2 Australia
503,,3
New Zealand Antler Growth5. 4.1
Introduction5.,4.,2
India and Sri Lanka5· h·3
Australia5.4.4
New Zealand37 37 38 39
4.0
l.i,.O
40 J.-5 45 45
1~5
46
l:-6
47 47 47
)+$
4S 4$ .
49
58 59
6o 60
61 61
6464
67 67
69
5.6
CHAPTER 6 6.1 , 6.2 6.3 6, I+
6.,5 606
6.7
6.8
CHAPTEH 7
PART IV
CHAPTER 8
Rate of Reproduction
5o5.1 Hind reproductive data Stog breedinB ag~
Nortality
BEHAVIOUfi
Introduction Animal Activity
Herd Size and Co~position
Tracking Behavio~r
Intraspecific and Interspecific J..ggression
Sexual Behaviour
6.6.,J.
6.6.2 6. 6.3
\'Jr:.11 ows
Rubbing Roaring Col!!rnunication
Trees
6.7.1 6.7.2 6.7.3
Auditory Visual Olfactory, Territoriality CHANI0v1ETHY Introductior- Skull Material Measurements 7 .3 .1 Crania
7.3.2 Antlers Age Deter~ination
7.4.1
Introduction 7.1+.2 MethodResults
7.,5.1
Cranial data7.5.2
Antler dataDISCUSSION AND CONCLUSIONS
72 72 72 '72
r1 r ( IJ~
74 75
77
78
?f;
78 BO 82 82
ct;
83 84
$6
86
90
9090 92 92
93 95 95
99
101 101
e.1
8.2 8.3
8.6 APPENDICES REFERENCES
H.equircrr.ent for Cover
Discussion of Forage Quality and Diet Selection
Disct~.ssion of Breeding and Antler Cycles
8.3.1 Australian breeding data 8.3 .2 1'Tew Zealand breeding data
BeJ.,J Antler cycling data Discussion of Craniometry 8.4.1 Cranial data
.<\.ntler data
Lntler defon~ities
Cor.irnents on the Future · of Sar.ibar in
New Zealand Conclusions
.101 102
107 107 109 111
113
113 113 114
115 118 121
134
I
I I I I I
IV
. v
VI VII
VIII IX
x
XI XII XIIILIST OF TABLES
Classification of sambar deer according to Wh:Ltchend (1972)
Previous works on dietary analysis Australian breeding data (a)c Based on
conception dates of 21 pregnant hinds (from Draisma, 1979)
Australian breeding data (b). Based on ages of 6o sambar not more than JO months
of age (Draisma, 1979)
Breeding data of New Zealand sambar Early antler grov..rth in sambar stags
(Draisma~ 1979)
Monthly variation of antler condition in Australian sambar. (Combined data from Bentley, 1978; and Draisma, 1979)
Monthlv variation of antler condition in New Ze~land sambar
Cranial measurements (Batcheler and McLennan, 1977)
Data analysis - Manawatu skulls Data analysis - New Zealand antlers
Forage species predominating in the sununer and winter diet of Moutoa deer
Means of cuticle area in faeces collected in April, August, November and February, and of individual species over ·the four combined collection dates
XIV Correlation matrix - Degree ..of correlation between cuticle area, fibre, energy, nitrogen
6 34
62
63 65 6S
70
71
91 98100 103
105
and water content 106
FIGURE
---
1 2 3
4
5 6 7
9
10
11
12
13
1.4
15
LIST OF FIGURES
The world \'lide distribution of sambar
as published by Whitehead (1972)
Distribution of sarnbar in the Manawatu (from Harris, 1966)
Distribution of sambar in the Bay of Plenty (from Harris, 1966)
Location of Moutoa
Map of reserve areas at Moutoa The· main study area
Monthly variation in the four major cuticle components of faeces
Monthly variation in fiv~ minor cuticle components of faeces
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in floating sweet grass
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in reed canarygrass
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in ryegrass (H
1)
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in tall f escue
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in Yorkshire fog
Seasonal variat.ion in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in flax
Seasonal variation in the content of Acid Detergent Fibre, Energy, Nitrogen and Water in poplar
Paue
. . ....
~'7 9
10 l.2
14
1'7 42
50
51
52
53
54
55
56
LIST OF FIGURES
16 Chart of tooth eruption in sambar
(fro~ Draisma, 1979)
17 Relationship between antler condition rutting activity in Australia .
18 Relationship between antler condition rutting activity in New Zealand
~
91+
and 108
and
110
PLATE
- ·- ·-
Frontispiece
1 2
3 4 5 6 7 8 9
10 11 12 1.3 14 15 16 17 lg 19
20
LIST OF PLATES
. _...,.._ ___
Sambar hindv Moutoa
(Phot ograph by Lo Barnard)
.Head of dead sambar hind
Area of medj_um density flax and unimproved pasture
Area of high density flax
Fully grown hind in dense flax cover (Photograph by L. Barnard)
Maximum height of tall f escue 'Massey Hide'
'Coffin Hide' Tree 'hide'
(Photograph by I. McKelvey) Evidence of feeding on flax Stag in poplar nursery
Sambar damage to young poplar bark Sambar hoof print
Cropped clump of tall f escue Moutoa stag in edge of clover/
ryegrass paddock · Skeleton of adult sambar hind
Small herd composed of two family groups (Photograph by Lo Barnard)
Typical sambar wallow 'Rubbing tree'
Detail of damage caused by antler rubbing
Head of typical sambar phenotype from Bay of Plenty
i i 2
15
15
16
20 22 22
23 26 26
27 27 29 31 66 76 79 81
PLATE 21 22
23
LIST OF' PLATF.S
Possj_ blc sambar / ).''·,1,r;a hyb:cid Typical rusa phenotype
Cementum layers in sectioned molar
·tooth
P~&r:.
