Advances in
Deer Biology
Praha 2006
Ludฤk Bartoลก, Adam Duลกek, Radim Kotrba,
Jitka Bartoลกovรก-Vรญchovรก (Editors)
Advances in
Deer Biology
Deer in a Changing World
Ludฤk Bartoลก, Adam Duลกek, Radim Kotrba,
Jitka Bartoลกovรก-Vรญchovรก (Editors)
Proceedings of the
6
th
International Deer Biology Congress
Prague, Czech Republic
7-11 August 2006
Advances in Deer Biology: Deer in a Changing World
Proceedings of the 6th International Deer Biology Congress,
Prague, Czech Republic, 7-11 August 2006
Editors:
Ludฤk Bartoลก, Adam Duลกek, Radim Kotrba,
Jitka Bartoลกovรก-Vรญchovรก
Ethology Group, Research Institute of Animal Production, Praha 10-Uhลรญnฤves, Czech
Republic (http://www.vuzv.cz/etolog/eetolog.htm)
Congress organised by
Research Institue of Animal Production, Prague
Czech University of Agriculture, Prague
Federation of European Deer Farmers Associations
and hosted by the Faculty of Agrobiology, Food and Natural Resources,
Czech University of Agriculture, Prague
Front cover photo and graphics Ludฤk Bartoลก
ยฉ 2006 Research Institute of Animal Production,
Praha, Czech Republic
Printed by PowerPrint, Provozovna - ฤZU,
Kamรฝckรก 129, 165 00 Praha 6 - Suchdol
ISBN 80-86454-73-8
Preface
This year's congress is a continuation in the tradition of previous congresses. The
"ancestor" to the International Deer Biology Congress series was the conference on
"Antler Development in Cervidae", organized in 1982 in Kingsville, Texas by Robert
Brown. This initial conference subsequently gave rise to the First International Deer
Biology Congress (1983, "Biology of Deer Production), organized by Ken Drew and
Peter Fennessy in Dunedin, New Zealand. Subsequent congresses were organized by
Bob Brown at Mississippi State, U.S.A. (1990), John Milne in Edinburgh, U.K.
(1994), Lรกszlรณ Sugรกr and Zoltรกn Zomborszky in Kaposvรกr, Hungary (1998), and
Michele Crรชte in Quรฉbec City, Canada (2002).
In the earlier congresses, complete papers were published in the Proceedings.
However, during later years the proportion of full papers declined reflecting an
increasing tendency in the academic world to present papers at conferences, but to
publish the full paper in an impacted journal afterwards. As a result, in the Fifth
International Deer Biology Congress, only a Scientific Program and Abstracts were
released. In the current Congress, we stepped back and present a compromise. While
this Proceedings consists predominantly of abstracts presented at the Congress, it also
contains longer contributions.
This Congress brought together a diverse group of deer scientists and deer
management professionals from Europe, North and South America, Asia, Australia
and New Zealand. Topics covered include deer management, diseases of deer,
genetics and evolution, management of endangered deer, reproduction, behaviour and
welfare, censusing and modelling populations, antler biology, responses of deer to
global environmental change, problems of deer overabundance, conservation of free
ranging populations, feeding ecology, seasonal and non-seasonal deer: (Arctic to
Tropic), venison and its potential contribution to human diet, and deer zooarcheology
and history. Thus the Advances in Deer Biology: Deer in a Changing World is a
comprehensive and up-to-date source of information on all current areas of deer
research.
Welcome to Prague, whether you are coming for the 6
th
International Deer Biology
Congress or just for a visit!
Ludฤk Bartoลก
Local Organizing Committee:
Ludฤk Bartoลก (Chairman),
Pavel ล ustr (www pages),
Radim Kotrba,
Jitka Bartoลกovรก-Vรญchovรก,
Radka ล รกrovรก (Excursions and Post-Congress tours),
Jan Pluhรกฤek,
Jorga Drรกbkovรก,
Josรฉ Panamรก,
John Fletcher (FEDFA);
+ unlimited number of other helpers
Scientific Steering Committee:
Jo Anne Smith-Flueck (Chairperson; Argentina)
Geoff Asher (New Zealand),
George Bubenik (Canada),
Norma Chapman (UK),
Matthew Cronin (USA),
John Fletcher (UK),
Susana Gonzรกlez (Uruguay),
Jerry Haigh (Canada),
Horst Kierdorf (Germany),
Afifullah Khan (India),
Karl Miller (USA),
Lรกszlรณ Sugรกr (Hungary),
Jimmy Suttie (New Zealand),
Nicholas Tyler (Norway).
Congress Secretary:
Ing. Martin Ledvinka
Unico Agric, Czech University of Agriculture Prague
Kamรฝckรก 129
165 21 Praha 6 โ Suchdol, Czech Republic
Contents
Plenary lectures .................................................. 1
1Priorities in Cervid conservation: Why science, zoogeography and
history do matter.
V. Geist ................................................ 2
2Deer responses to global environmental changes.
M. C. Forchhammer ....................................... 2
3 Emerging disease in wild and captive Cervids.
M. R. Woodbury and J. R. Campbell .......................... 4
4 Seasonal versus non-seasonal reproduction in deer: From the arctic
to the tropics.
G. A. Bubenik .......................................... 16
5Fallow deer, lekking and alternative mating strategies in San Rossore,
Italy: insights from a long term study.
M. Apollonio ........................................... 17
6Recent progress in antler regeneration and stem cell research.
C. Li and J. M. Suttie ..................................... 18
7Conservation of tropical deer: what does the future hold?
W. J. McShea ........................................... 19
Deer management ............................................... 21
8Three years of roe deer (Capreolus capreolus) radio-tracking in a
Mediterranean environment.
A. J. Ferreira and C. Silva ................................. 22
9Seasonal home range shift of red deer in a forest-agriculture area,
Hungary.
L. Szemethy, Zs. Birรณ, K. Katona, K. Mรกtrai, Sz. Orosz, and N. Bleier
...................................................... 22
10 Space use patterns of Persian fallow deer following reintroduction.
S. Bar-David, D. Saltz, A. Dolev, A. Perelberg, and T. Dayan ..... 23
11 Population size and demographic variables of red deer in Bydgoszcz
National Forest, central Poland.
P. Beszterda ............................................ 24
12 Management of red deer in Poland: field data versus official hunting
statistic.
B. Bobek, T. Mamok, J. Mikoล, W. Rembacz, A. Standio, and R.
Wasilewski ............................................ 24
13 Over-abundance of deer: Is shooting the answer?
D. C. MacMillan ........................................ 25
14 Slaughter records as a body condition indicator or reindeer - How can
records be improved?
A. Olofsson, B. ร
hman, and ร. Danell....................... 26
15 The management of reindeer in the Mongolian Tsaatan culture.
J. C. Haigh and M. G. Keay ............................... 29
16 Can supplementary feeding improve productivity in reindeer
husbandry?
B. ร
hman and ร. Danell .................................. 30
17 Twenty years of impact of the Chernobyl accident on reindeer
management and meat production in Sweden and Norway.
B. ร
hman and L. Skuterud ................................ 32
18 Economic sustainability of farmed venison production in the UK.
M. H. Davies and D. G. Chapple............................ 35
19 Public perception of deer management and control strategies.
M. I. Malins............................................ 36
20 Preferences of red deer for subtropical pasture species.
G. M. Dryden and K. J. Whelan ............................ 37
21 Evaluation of forage herbs for farmed red deer: feeding value and
trace elements.
S. O. Hoskin, P. R. Wilson, M. Ondris, and A.-H. Bunod ........ 38
22 Habitat Utilization by Himalayan Musk Deer (Moschus chrysogaster),
Sambar ( Cervus unicolor) and Barking Deer (Munitacus muntjac) at
Kedarnath Wildlife Sanctuary, Western Himalaya.
S. Sathyakumar, S. N. Prasad, G. S. Rawat, and A. J. T. Johnsingh . 39
23 Feeding habits of red deer in Hungarian forested and agricultural
areas.
K. Katona, L. Szemethy, K. Mรกtrai, N. Bleier, and Sz. Orosz ..... 40
24 Detection of needles: tool for evaluation of diet quality in wild
ruminants.
J. Kamler, M. Homolka, M. Heroldovรก, M. Baranฤekovรก, and J. Prokeลกovรก
...................................................... 41
25 Environmental factors affecting Scots pine debarking by red deer in
south-western Poland.
J. Borkowski and P. Nasiadka .............................. 41
26 New technique for estimation Cervidea hiding cover.
A. J. Ferreira and A. M. Oliveira ............................ 42
27 Calving sites fidelity in free-ranging moose.
J.-P. Tremblay, E. Solberg, B.-E. Sรฆther, and M. Heim .......... 43
28 Spatio-temporal distribution of white-tailed deer relative to prescribed
burns on rangeland in south Texas, USA.
M. G. Meek, S. M. Cooper, M. K. Owens, and A. L. Wappel ...... 44
29 Sexual segregation and differences in quality of diet in white-tailed
deer (Odocoileus viginianus mexicanus) in a tropical dry forest in
Mexico.
A. Buenrostro, S. Gallina, and G. Sรกnchez-Rojas ............... 45
30 Sex comparison of linear body measures of growing red deer calves
(Cervus elaphus hippelaphus).
B. Dmuchowski, M. Snochowski, and A. Krzywiลski............ 46
31 The influence of management system of farmed fallow Deer (Dama
dama) on selected production traits during winter season.
B. Dmuchowski, J. Starz, A. Demiaszkiewicz, and R. Niลผnikowski
...................................................... 47
32 Deer home range overlap and habitat heterogeneity in Northeastern
Mexico.
J. Bello, S. Gallina, M. Equihua, and N. Corona ................ 47
33 Influence of ranging strategy on home range size: red deer hinds in a
forest-agriculture habitat.
Zs. Birรณ, L. Szemethy, and K. Katona ........................ 48
34 New project on red deer Cervus elaphus in Sweden.
A. Jarnemo ............................................. 49
35 Mapping of male red deer Cervus elaphus movements in southern
Sweden.
A. Jarnemo ............................................. 49
36 Importance of floodplain forest for deer management.
J. Prokeลกovรก, M. Baranฤekovรก, and M. Homolka ............... 50
37 Gross composition and protein fractions of milk from fallow deer
(Dama dama).
G. M. Pisani, M. Malacarne, C. S. Soffiantini, P. Franceschi, P.
Formaggioni, E. Piasentier, A. Summer, and P. Mariani.......... 51
38 Estimating in vitro digestibility of wild sika deer (Cervus nippon
yesoensis) in Hokkaido, Japan.
C. Yayota, K. Nishitani, K. Ueda, Y. Yanagawa, Y. Matsuura, M. Suzuki,
H. Hata, and S. Kondo .................................... 52
39 Comparison of physical condition of two Red deer (Cervus elaphus)
populations.
A. J. Ferreira and R. M. Ramalho ........................... 53
40 Distribution, abundance and management of the two native deer in
Italy.
L. Carnevali, F. Riga, and S. Toso .......................... 53
41 Interspecific competition between large herbivores: the fallow deer -
roe deer case.
P. Kjellander ........................................... 54
42 Current knowledge of the Central American red brocket deer
(Mazama temama Kerr, 1792) in Mexico.
J. Bello-Gutiรฉrrez ....................................... 55
43 Energy requirement of captive grey brocket deer (Mazama
gouazoubira) determined by weight equilibrium and double-labeled
water.
A. Berndt, M. Z. Moreira, J. M. B. Duarte
, J. Barbosa, and D. P. D. Lanna
...................................................... 56
44 Modelling the influence of resources on the distribution and
aggregation of red deer hinds during the rut: implications for mating
system and management.
J. Pรฉrez-Gonzรกlez, A. M. Barbosa, and J. Carranza ............. 56
45 Red deer as a newcomer in Estonian fauna.
T. Randveer and E. Niittee ................................ 57
46 Comparison of different weaning times of farmed Hungarian red deer
(Cervus elaphus hippelaphus) calves.
Z. Pados, J. Szabรณ, J. Nagy, Sz. Nagy, and Z. Zomborszky ....... 58
47 A photographic guide for aging fallow deer Dama dama.
A. M. De Marinis, C. Gozzi, V. Marasco, and S. Toso........... 59
Diseases of deer ................................................. 61
48 Recent advances in health and welfare of farmed deer in New
Zealand.
P. R. Wilson ............................................ 62
49 Health and production challenges facing intensive deer farming
industries.
P. R. Wilson ............................................ 63
50 Chronic wasting disease in North America - A deer farmerโs
perspective.
C. Tedford ............................................. 64
51 Chronic wasting disease in Canadian wildlife: An expert opinion on
the epidemiology and risks to wild deer .
C. Maxwell............................................. 65
52 Epidemiological investigations of Johne's disease in deer.
J. C. Glossop, P. R. Wilson, C. Heuer, and G. Nugent ........... 66
53 Johne's disease in farmed deer in New Zealand.
C. G. Mackintosh, J. F. T. Griffin, and G. W. de Lisle ........... 67
54 Insights into the pathogenesis of Johneโs disease in red deer (Cervus
elaphus).
C. G. Mackintosh, J. Thompson, J. F. T. Griffin, and G. W. de Lisle
...................................................... 68
55 The efficacy of oral and pour-on ivermectin and pour-on moxidectin
in farmed red deer.
S. O. Hoskin, W. E. Pomroy, P. R. Wilson, M. Ondris, and P. Mason
...................................................... 69
56 An internatinal review of Leptospirosis in wild and farmed deer.
M. A. Ayanegui-Alcรฉrreca, P. R. Wilson, C. Heuer, J. M. Collins-
Emerson, C. G. Mackintosh, A. C. Midwinter, and F. Castillo-Alcala
...................................................... 70
57 Epidemiology of Leptospiral infections with Serovars Hardjobovis,
Pomona and Copenhageni in farmed red deer (Cervus elaphus) in New
Zealand.
M. A. Ayanegui-Alcรฉrreca, P. R. Wilson, C. G. Mackintosh, J. M. Collins-
Emerson, C. Heuer, A. C. Midwinter, and F. Castillo-Alcala
...................................................... 71
58 Anthelmintic use and internal parasite control in farmed deer in New
Zealand.
F. Castillo-Alcala, P. R. Wilson, W. E. Pomroy, and S. O. Hoskin . 72
59 Subdural occurrence of Elaphostrongylus cervi and Setaria cervi in red
deer of West Hungary.
L. Sugรกr, Sz. Kovรกcs, and A. Kovรกcs ........................ 73
60 Disease problems in Mongolian reindeer.
J. C. Haigh, M. G. Keay, V. Gerwing, J. Erdenbaatar, and M. Nansalmaa
...................................................... 73
61 Histopathology of fluorotic coronal dentine of roe deer (Capreolus
capreolus) and red deer (Cervus elaphus) teeth.
H. Richter, A. Richards, and H. Kierdorf ..................... 74
62 Mineral composition and requirements for growth of farmed red deer
in New Zealand.
F. Castillo-Alcala, P. R. Wilson, and N. D. Grace .............. 75
63 Recent advances in understanding therapy with Copper Oxide Wire
Particles in New Zealand Farmed deer.
P. R. Wilson, F. Castillo-Alcala, and N. D. Grace .............. 76
64 Nasopharyngeal bot fly, Oestridae larvae in red deer in Hungary.
L. Sugรกr, Sz. Kovรกcs, and A. Kovรกcs ........................ 77
65 ITS2 sequences of Dictyocaulus lungworms from red and fallow deer
in Hungary: molecular evidence for a new genotype.
Z. รcs, L. Sugรกr, and Z. Pรฉnzes ............................ 78
66 Fascioloidosis of red deer and its therapy in ''Szigetkรถz'' region in the
North-West of Hungary (1998-2005).
B. Egri and E. Giczi...................................... 78
67 Coprological monitoring of Trematodes in free-ranging red deer
population at eastern Croatia.
A. Slavica, T. Florijanฤiฤ, Z. Janicki, D. Konjeviฤ, K. Severin, R. Beck
and K. Pintur ........................................... 79
68 Sub-clinical parasitism, weaning date, growth of deer fawns and
reproductive performance of hinds .
J. M. Mwendwa, M. L. W. J. Broekhuijse, S. O. Hoskin, W. E. Pomroy,
and P. R. Wilson ........................................ 80
69 Investigation of the sanitary status of red deer (Cervus elaphus) culled
in the Italian Alps between 2001 and 2005.
E. Andreoli, I. Bertoletti, A. Bianchi, E. Heinzl, E. Scanziani, and S.
Mattiello .............................................. 81
70 General comparison of taxonomic characters distinguishing two
closely related species of deer lice - Solenopotes burmeisteri and S.
capreoli (Phthiraptera, Linognathidae).
V. Bรกdr, P. ล tindl, and J. Preisler............................ 82
Genetics and Evolution ........................................... 83
71 Landscape features affect gene flow of Scottish Highland red deer
(Cervus elaphus).
S. Perez-Espona, J. McLeod, F. J. Perez-Barberia, C. G. I. Jiggins, and J.
Pemberton ............................................. 84
72 Sex biased dispersal in an expanding red deer population.
H. Haanes, K. H. Rรธed, and O. Rosef ........................ 84
74 A molecular phylogeny of the evolutionary radiation of New World
deer (Odocoileinae, Cervidae): Implications for biogeography and the
evolution of antlers.
S. M. Carr, E. D. Richards, H. D. Marshall, and J. M. Smith-Flueck
...................................................... 85
75 Genetic distinctiveness of isolated and threaten Tsaatan reindeer
herds in Mongolia.
K. H. Rรธed, J. C. Haigh, V. Gerwing, and M. Keay ............. 86
76 Conservation genetics of Argentinean pampas deer populations.
S. Gonzรกlez, M. Cosse, V. Raimondi, M. L. Merino, B. Galvan, and J. E.
Maldonado ............................................. 87
77 Genetic characterisation of roe deer (Capreolus capreolus) population
of Parma Apennines.
C. S. Soffiantini, G. M. Pisani, M. Malacarne, G. Gandolfi, A. Sabbioni,
and J. Tagliavini......................................... 88
78 Aplotypic characterization of roe deer by asymmetric PCR and SSCP
analysis.
J. Tagliavini, S. Casagrande, M. Malacarne, and P. Mariani ....... 88
79 Phylogeography of Iberian red deer populations and their
relationships with main European red deer lineages.
J. L. Fernรกndez-Garcรญa, J. G. Martรญnez, L. Castillo, and J. Carranza
...................................................... 89
80 The artificial occurrence of the fallow deer, Dama dama dama (L.,
1758), on the island of Rhodes (Dodecanese, Greece): insight from
mtDNA analysis.
M. Masseti, A. Cavallaro, E. Pecchioli, and C. Vernesi ........... 90
81 Comparative anatomy of three Asian ruminant animals.
J. Kimura and K. Fukuta .................................. 91
82 Characterization of the growth curve of red deer (Cervus elaphus
scoticus) in a herd in Central Mexico.
A. C. Delgadillo, R. Lรณpez, H. H. Montaldo, J. M. Berruecos, A. Luna,
and G. C. Vรกsquez....................................... 92
83 Mitochondrial DNA variability and polymorphism of ISSR-PCR
markers in the reindeer population of Eastern Siberia.
N. V. Kol, O. E. Lazebny, and I. A. Zakharov ................. 95
84 A new conservation genetic union from Pampas deer (Ozotoceros
bezoarticus) in Southern Brazil.
F. G. Braga, S. Gonzรกlez, and J. E. Maldonado ................ 96
85 DNA microsatellite analysis for parentage control of red deer in Czech
Republic.
M. Ernst............................................... 97
Management of endangered deer ................................... 99
86 Status, ecology and conservation of barasingha (Cervus duvauceli
duvauceli) in Terai grasslands of Northern India.
J. A. Khan and A. Kaleem................................ 100
87 Swamp deer in Uttaranchal state, India.
S. P. Sinha, S. Chandola, and B. C. Sinha .................... 101
89 Swamp deer (Cervus duvaceli) habitat evaluation using remote sensing
and GIS in Suklaphanta Wildlife Reserve, Nepal.
T. B. Thapa ........................................... 103
90 Population Ecology of Hangul (Cervus elaphus hanglu) in Dachigam
National Park, Kashmir, India.
A. Khursheed, S. Sathyakumar, and Q. Qureshi ............... 104
91 Microsatellite variation of Hainan Eld-s deer (Cervus eldi hainanus)
in China: Implications for conservation.
Q. Zhang, Y.-L. Song, D.-X. Zhang, and Z. Zeng .............. 105
92 Social structure of the reintroduced Persian fallow deer (Dama
mesopotamica) population: integrating three observation methods.
A. Perelberg, S. Bar-David, U. Roll, A. Dolev, and D. Saltz ..... 105
93 Ecology and conservation of the huemul in southern Chile.
R. Gill, C. Saucedo, and D. Aldridge ....................... 106
94 Status, genetic structure and Conservation suggestion of Chinese
water deer.
M. Chen and E. Zhang ................................... 107
95 Spatial pattern characteristics of wapiti habitat fragmentation factors
based on spatial autocorrelation and semi-variance analysis in
Northeastern China.
M. Zhang, G. Jiang, and J. Ma............................. 108
96 Assisted reproductive technologies for endangered deer species.
Y. Locatelli, J.-C. Vallet, X. Legendre, and P. Mermillod ........ 109
97 Diet composition and habitat selection of red deer during winter in
Helan Mountains, China.
Z. S. Liu and X. M. Wang ................................ 110
98 Conservation status quo and study progress of Siberian musk deer
(Moschus moschiferus) in China.
J. Wu and Y. Zhang ..................................... 111
99 Agonistic and non-agonistic behaviour interactions in Indian
blackbuck (Antelope cervicapra L.) during dominance hierarchy
formation.
T. Rajagopal and G. Archunan ............................. 111
Reproduction .................................................. 113
100 Gossypol-based contraception in male deer (Cervus elaphus).
Z. Giลผejewski, B. Szafranska, Z. Steplewski, G. Panasiewicz, and H.
Koprowski ............................................ 114
101 The hoarse vocalization and the inflatable laryngeal air sac of reindeer
(Rangifer tarandus).
R. Frey, A. Gebler, G. Fritsch, K. Nygrรฉn, and G. E. Weissengruber
..................................................... 114
102 Patterns of long-term reproductive success in male and female white-
tailed deer.
R. W. DeYoung, K. L. Gee, S. Demarais, R. L. Honeycutt, and R. A.
Gonzales.............................................. 115
103 Observations on the reproductive behaviour of sambar deer (Cervus
unicolor unicolor) in a bush enclosure in Victoria, Australia.
W. M. Harrison, I. A. Moore, M. Draisma, and G. I. Moore ...... 116
104 Sexual choice in lekking fallow deer (Dama dama): variable female
strategies.
S. Imperio, S. Focardi, F. Ronchi, and A. M. De Marinis ........ 117
105 Variation in fawn production in a semi arid environment: An
energetics approach.
D. G. Hewitt and E. L. Monaco ............................ 118
106 Movements of female white-tailed deer during parturition and the rut
in a high-quality, balanced sex ratio herd in Maryland, USA.
L. I. Muller, K. A. Adams, M. C. Conner, and J. L. Bowman ..... 119
107 Refrigerated storage impairs chromatin of Iberian red deer (Cervus
elaphus hispanicus) epididymal spermatozoa kept inside the
epididymis.
A. E. Dominguez-Rebolledo, M. C. Esteso, M. R. Fรฉrnandez-Santos, D.
Matias, F. Martinez-Pastor, and J. J. Garde................... 120
108 Immunohistochemical expression of steroidogenic enzymes in the
corpus luteum and placenta of sika deer (Cervus nippon) during
pregnancy.
Y. Matsuura, D. Hayakawa, Y. Yanagawa, M. Sasaki, H. Igota, C. Yayota,
S. Kondo, N. Kitamura, T. Tsubota, and M. Suzuki ............ 121
109 Objective quality control of frozen-thawed red deer spermatozoa by
Computer-Assisted Semen Analysis - instrument settings.
Sz. Nagy, E. Puskรกs, I. Pรฉntek, and Z. Zomborszky ............ 122
110 Immunohistochemical expression of androgen receptor (AR), estrogen
receptor alpha (ER) and estrogen receptor beta (ER) in the caudal
and metatarsal glands of sika deer (Cervus nippon).
M. Suzuki, Y. Yanagawa, Y. Matsuura, S. Otsuka, D. Hayakawa, M.
Sasaki, C. Yayota, H. Igota, S. Kondo, and N. Kitamura ........ 123
111 Comparison of estrogen receptor and progesterone receptor
expression during the estrus and pregnancy in uteri of sika deer
(Cervus nippon).
Y. Yanagawa, Y. Matsuura, D. Hayakawa, C. Yayota, M. Sasaki, S.
Kondo, N. Kitamura, and M. Suzuki ........................ 124
112 Roaring trends in red deer: a preliminary analysis.
A. Bocci, K. Attinault, and M. Telford ...................... 125
Behaviour and welfare .......................................... 127
113 Assessing the performance of a Persian fallow deer population 10
years after reintroduction.
D. Saltz and S. Bar-David ................................ 128
114 Social competence in Chinese muntjac deer.
A. Fischer and H. Hendrichs .............................. 129
115 The analysis of sexual segregation in fallow deer (Dama dama) on
different time and space scales.
S. Ciuti, S. Luccarini, and M. Apollonio ..................... 130
116 Behavioural modifications of female ungulates during late pregnancy
and early lactation: the case of fallow deer Dama dama.
S. Grignolio, P. Bongi, S. Ciuti, E. Bertolotto, and M. Apollonio . . 131
117 Pre-orbital gland opening in red deer (Cervus elaphus) calves: Signal
of excitement?
J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, and L. ล vecovรก .............. 132
118 The effect of the birth weight on the calfโs allosucking success in the
red deer (Cervus elaphus) supports the compensation hypothesis.
A. Duลกek
.
and L. Bartoลก .................................. 132
119 Do red deer (Cervus elaphus) grandmothers nurse their
grandchildren?
J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, J. Drรกbkovรก, L. ล vecovรก, J. Pluhรกฤek, R.
Kotrba, A. Duลกek ....................................... 133
120 When prey fight back: higher levels of aggressive defence by mule
deer than whitetail females lowers vulnerability of mule deer fawns to
coyotes early in life.
S. Lingle, W. F. Wilson, and S. M. Pellis .................... 134
121 Why Help? The evolution of altruistic antipredator defence in mule
deer.
S. Lingle, D. Rendall and S. M. Pellis ....................... 135
122 Cooperative anti-predatory behaviour in sympatric white-tailed,
fallow, roe and red deer: Experimental confirmation using a dummy.
R. Kotrba, L. Bartoลก, J. Bartoลกovรก-Vรญchovรก, J. Panamรก, V. Kลกรกda, P. ล ustr,
J. Pluhรกฤek, A. Duลกek, D. Vaลkovรก-Formanovรก, G. Illmann, E. ล mรญdovรก,
and K. V. Miller
........................................ 136
123 Rutting encounter between males and female choice in fallow deer
(Dama dama).
B. Friฤovรก, L. Bartoลก, J. Bartoลกovรก-Vรญchovรก, J. Panamรก, P. ล ustr, and E.
ล mรญdovรก .............................................. 137
124 Habitat selection and home range size of red deer (Cervus elaphus) in
montane areas of ล umava National Park, Czech republic -
preliminary results.
P. ล ustr and A. Jirsa ..................................... 138
125 Sex-specific strategies of dentine depletion in red deer.
J. Carranza, C. Mateos, S. Alarcos, C. B. Sรกnchez-Prieto, and J. Valencia
..................................................... 138
126 Does a hindโs rank affect duration of filial and non-filial calfโs nursing
in red deer (Cervus elaphus)?
J. Drรกbkovรก, J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, J. Pluhรกฤek, R. Kotrba, L.
ล vecovรก, and A. Duลกek .................................. 139
127 ISAMUD: an integrated software environment for analysis and
management of GPS telemetry data.
F. Cagnacci, F. Urbano, C. Furlanello, M. Neteler, and L. Pedrotti
..................................................... 140
Censusing and modelling populations .............................. 143
128 Censusing and modelling of red deer (Cervus elaphus L.) populations
in Poland by using โInventโ and โAntler-2000โ software.
B. Bobek, W. Frฤ
ckowiak, M. Gawor, M. Kolecki, D. Merta, and L.
Wiลniowska........................................... 144
129 Estimating Red deer populations abundance in the Alps: successful
experiments on night surveys.
S. Focardi, B. Franzetti, A. Monaco, and L. Pedrotti ........... 145
130 Whitetailed Deer Density Estimation Using Thermal Infrared
Imaging.
P. A. Tappe and R. E. Kissell ............................. 146
131 Estimating red deer Cervus elaphus populations: an analysis of
variation and cost effectiveness of counting methods.
M. J. Daniels .......................................... 151
132 Simple Movement Models for Complex Animals in Heterogeneous
Landscapes.
J. M. Morales.......................................... 152
133 Reconstruction of the male population of red deer in Hungary.
S. Csรกnyi ............................................. 152
134 The second mass-mortality of an introduced sika deer population.
H. Takahashi and K. Kaji ................................ 153
135 Fecal-pellet group count as index of sika-deer (Cervus nippon)
population density on subalpine plateau in Japan.
R. Goda, M. Ando, H. Sato, and E. Shibata .................. 154
136 Comparison of four techniques to estimate roe deer abundance in
Alpine areas.
N. Putzu, V. La Morgia, and F. Bona ....................... 154
137 Distance sampling and pellet group count to assess deer populations:
an application to conservation and management in the Alps.
L. Pedrotti, F. Cagnacci, I. Callovi and A. Tagliabรฒ ............ 155
138 Red deer (Cervus elaphus) space use and population dynamics in two
Alpine National Parks.
F. Filli, L. Pedrotti, and H. Gunsch ......................... 156
139 A population-dynamic study of red deer in Baranya, Somogy, Tolna
and Zala counties from 1970 to 2006.
R. Barna and L. Sugรกr ................................... 157
Antler biology .................................................. 159
140 Visualization and characterization of stem cells from the regenerating
deer antler.
H. J. Rolf, U. Kierdorf, H. Kierdorf, N. Seymour, J. Napp, H. Schliephake,
and K. G. Wiese........................................ 160
141 Antlers may regenerate from persistent neural crestlike stem cells.
J. G. Mount, M. Muzylak, S. Allen, S. Okushima, T. Althnaian, I. M.
McGonnell, and J. S. Price ................................ 161
142 Stem cells isolated from the regenerating antler express key markers
of the osteogenic lineage.
J. Napp, K. G. Wiese, U. Kierdorf, H. Kierdorf, N. Seymour, H.
Schliephake, and H. J. Rolf ............................... 162
143 Mitogenic effects of androgens on mixed antler cell cultures.
H. J. Rolf, K. G. Wiese, G. A. Bubenik, L. Bartoลก, R. Kotrba, I. Lรผtjens
and H. Schliephake...................................... 163
144 Antler growth in red deer stags (Cervus elaphus) depends on
testosterone, but not IGF-1, LH, prolactin or cortisol.
L. Bartoลก, D. Schams, J. ล iler, S. Losos, and G. A. Bubenik...... 164
145 Fetal differentiation of the antler developing area in red deer (C.
elaphus).
P. M. F. Audenaerde and P. J. M. Simoens ................... 164
146 Central vessels in roe deer antlers (Capreolus capreolus) - a
histomorphological study.
H. J. Rolf and C. H. Lohmann............................. 165
147 Antler characteristics of the Sardinian red deer (Cervus elaphus
corsicanus): a preliminary analysis.
A. Caboni, C. Murgia, and S. Mattioli ....................... 166
148 What we can learn from antler composition and structure: from
nutrition to management.
T. Landete-Castillejos, J. A. Estรฉvez, A. J. Garcia, F. Ceacero, E. Gaspar-
Lรณpez, D. Carriรณn, and L. Gallego ......................... 167
149 Post-velvet shedding antler histology of red deer (Cervus elaphus)
living in the wild.
A. Dobrowolska and K. Gรณrecka .......................... 168
150 Lengths of pedicles and antlers in Reeves' muntjac.
N. G. Chapman ........................................ 169
151 Consistent interindividual variability in proliferation potential of
antler cells cultivated in vitro under various treatments.
E. Kuลพmovรก
,
, L. Bartoลก, M. Tomรกnek, R. Kotrba, and G. A. Bubenik
..................................................... 169
Responses of deer to global environmental change .................... 171
152 Biogeography of Cervidae in Peru.
J. Barrio.............................................. 172
153 The influence of season, food intake, and social rank on cortisol
secretion in red deer (Cervus elaphus).
F. Balfanz, C. Beiglbรถck, S. Huber, R. Palme, and W. Arnold.... 173
154 Defense of territories by rutting red deer stags, Cervus elaphus, in
Patagonia, Argentina.
J. M. Smith-Flueck and W. T. Flueck ....................... 174
155 Spatial behavior paths of food search in roe deer (Capreolus
capreolus).
S. Said, M. Pellerin, M. Le Corre, O. Widmer, and G. Van Laere . 178
156 Carbon and nitrogen efficiencies in venison production.
M. H. Davies, D. G. Chapple, and B. Cottrill ................. 179
157 Methane production by farmed red deer.
N. M. Swainson, S. O. Hoskin, and H. Clark ................. 180
158 Why the Patagonian huemul deer in Argentina fails to recover: An
ecological hypothesis.
W. T. Flueck and J. M. Smith-Flueck ....................... 181
159 Deer management and private hunting? Turning point for
management system in Japan.
A. Takayanagi ......................................... 186
160 The biology of antler growth in endangered Bawean deer (Axis kuhlii).
G. Semiadi, K. Subekti, B. Masyud, I. K. Sutama, and L. Affandy
..................................................... 187
161 The preservation of rusa stag semen using TRIS egg yolk diluent with
different carbohydrate and storage temperature.
W. M. M. Nalley , R. Handarini, G. Semiadi, M. R. Toelihere, T. L.
Yusuf, and B. Purwantara ................................ 187
162 Semen quality of rusa stags (Cervus timorensis) during one antler
cycle.
R. Handarini, W. M. M. Nalley, G. Semiadi, M. R. Toelihere, S.
Agungpriyono, B. Purwantara, and Subandriyo ................ 189
Problems of deer overabundance ................................... 191
163 A test of localized management in a white-tailed deer herd.
B. F. Miller, R. W. DeYoung, T. A. Campbell, B. R. Laseter, W. M. Ford,
and K. V. Miller ........................................ 192
164 Do wildlife warning reflectors alter white-tailed deer behavior along
roadways?
G. J. D'Angelo, J. G. D'Angelo, G. R. Gallagher, D. A. Osborn, K. V.
Miller, and R. J. Warren .................................. 193
165 Cascading effects of long term chronic browsing on lifehistory traits
in white-tailed deer.
S. D. Cรดtรฉ, A. Simard, R. B. Weladji, and J. Huot.............. 194
166 Regeneration dynamics of boreal forests along an experimental
gradient of deer densities.
J.-P. Tremblay, J. Huot, and F. Potvin ....................... 195
167 Impacts of cervids on invertebrate communities on forest floor in
relation to deer species, density and site productivity.
O. Suominen, I. L. Persson, and T. Saikkonen ................. 196
168 Sustainable population density of red deer in Mediterranean
ecosystems.
J. Carranza, J. Torres, S. Alarcos, J. Pรฉrez-Gonzรกlez, C. B. Sรกnchez-Prieto,
C. Mateos, L. Castillo, and J. Valencia ...................... 197
169 Influence of population density on white-tailed deer foraging behavior
and activity budget.
M.-L. Coulombe, S. D. Cรดtรฉ, and J. Huot .................... 198
170 Trade-off between food and cover: summer movements and activity
budget in white-tailed deer.
A. Massรฉ, S. D. Cรดtรฉ, and J. Huot .......................... 199
171 Relationships between moose (Alces alces) pellet groups and
characteristics of forests.
R. Heikkilรค ........................................... 200
Conservation of free ranging populations: conflicts of interest ........... 207
172 Status, distribution and conservation of musk deer (Moschus
chrysogator) in Kedarnath Wildlife Sanctuary, Uttranchal Himalayas,
India.
O. Ilyas .............................................. 208
173 Seed dispersal by the reintroduced Persian fallow deer in the Judean
Mountains, Israel.
R. Zidon, D. Saltz, and U. Motro .......................... 209
174 Deer management and monitoring of browsing impacts in Austrian
national parks.
R. Zink and F. Reimoser ................................. 210
175 Wildlife trade in deer species: A need for developing wildlife forensic
techniques.
S. P. Goyal, A. Mandal, R. R. Singh, S. Mishra, and C. P. Sharma
..................................................... 210
176 Habitat use of pampas deer (Ozotoceros bezoarticus) at agricultural
areas in southern Brazil.
F. G. Braga ........................................... 211
177 Impact of red deer browsing on the understory of Hungarian forests.
N. Bleier, K. Katona, L. Szemethy, J. Szรฉkely, M. Nyeste, ร. Fodor, A.
erhes, V. Kovรกcs, and T. Olajos ........................... 212
178 Effects of small barriers on habitat use in red deer.
C. B. Sรกnchez-Prieto
,
, J. Carranza, S. Alarcos, and C. Mateos .... 213
Feeding ecology ................................................ 215
179 Botanical composition of taruka (Hippocamelus antisensis) diet during
rainy season in Huascaran National Park, Peru.
C. Gazzolo ............................................ 216
180 Habitat use by two large deer species (Hippocamelus antisensis and
Odocoileus virginianus) and one small deer species (Mazama bricenii)
in the Apolobamba Integrated Management Natural Area (La Paz-
Bolivia). ............................................. 217
181 Impact of deer browsing and other environmental factors upon
growth and development of fir saplings (Abies alba Mill.) in the
Bieszczady Mountains, southern Poland.
D. Merta and K. Kumรณr .................................. 218
182 Why deer strip bark? -two case studies of bark stripping by sika deer
in central Japan.
M. Ando, Z. Jiang, and E. Shibata .......................... 219
183 Influence of an extreme climatic event on the winter diet of red and
roe deer in northeastern France.
D. B. F. Storms, S. Said, J.-L. Hamann, C. Saint-Andrieux, J.-L. Wilhelm,
and F. Klein ........................................... 220
Seasonal and non-seasonal deer: (Arctic to Tropic) .................... 221
184 Seasonal migration pattern of red deer ( Cervus elaphus L.) in the
central Slovakian mountains.
S. Finฤo, J. Buฤko, and S. Steyaert ......................... 222
185 Scale-dependent habitat selection of GPS-collared Alpine red deer the
role of food availability and quality.
B. Zweifel-Schielly and W. Suter .......................... 223
186 Photic modulation of the temporal pattern and rate of activity in
reindeer.
B. E. H. van Oort, N. J. C. Tyler, M. P. Gerkema, L. Folkow, and K. A.
Stokkan .............................................. 224
187 Habitat use and selection of fallow deer (Dama dama L.) in a
Mediterranean environment.
P. Di Luzio, P. Montanaro, and S. Focardi.................... 225
188 Function of habitat segregation in regulation of isolated sika deer
population.
S. Tatsuzawa .......................................... 226
Venison and its potential contribution to diet ......................... 227
189 Venison and the history of early European hunting enclosures.
T. J. Fletcher........................................... 228
190 Fatty acid profiles in Javan rusa (Cervus timorensis russa) stags.
R. Sookhareea, R. Tume, W. R. Shorthose, and G. M. Dryden .... 229
191 The effect of pelvic suspension on the biochemical and sesnory quality
of venison from red deer (Cervus elaphus) and fallow deer (Dama
dama).
C. L. Hutchison, J. S. Flesch, and R. C. Mulley................ 234
192 Contents of toxic metals (Cd, Pb, Hg) in tissues of the red deer (Cervus
elaphus) living in the wild.
A. Dobrowolska and K. Gรณrecka .......................... 240
193 Variations in characteristics of fat, free amino acids and taste of meat
of Japanese deer.
M. Ishida, T. Inoue, T. Mashiko, K. Souma, and S. Ikeda ....... 241
Deer zooarcheology and history ................................... 243
194 Stable isotopes evidence of seasonality effects on diet and locomotor
adaptations of Pleistocene deer from southern Spain.
J. A. Estรฉvez, A. Grandal-dโAnglade, T. Landete-Castillejos, A. J. Garcรญa,
and
L. Gallego ........................................ 244
195 Biometry and palaeoecology of the Red deer (Cervus elaphus Linnรฉ,
1758) during middle and upper Pleistocene in Western Europe. The
example of the Lazaret cave (Alpes-Maritimes; France)
M. Liouville, P. Valensi, and E. Psathi ...................... 248
196 Fallow deer of Rhodes: an ongoing, comprehensive study about
ecology, genetics and conservation.
D. Mertzanidou and A. Legakis ........................... 249
Contributions received and accepted after the deadline ................. 251
197 Conservation of huemul (Hippocamelus bisulcus) deer in Chilean
Patagonia: a new research initiative.
P. Corti .............................................. 252
198 Translocation and semi-captive breeding of huemul (Hippocamelus
bisulcus) with purpose of reintroduction in Chile.
P. Corti .............................................. 253
199 So similar and yet so different: The surprising polyphyletic origin the
genus Mazama (Mammalia: Cervidae).
J. M. B. Duarte, S. Gonzรกlez, and J. E. Maldonado ............ 254
200 Factors affecting the composition of autumn diet of red deer (Cervus
elaphus) in Alpine environment.
M. Heroldovรก, M. Homolka, J. Kamler, C. Ghezzi, W. Redaelli, E.
Andreoli, and S. Mattiello ................................ 255
Author index .................................................. 257
Index ........................................................ 263
Plenary lectures
2 R
1
Priorities in Cervid conservation: Why science, zoogeography and
history do matter.
V. Geist
Professor Emeritus of Environmental Science, The University of Calgary, Calgary,
Alberta, Canada
A review of the long history of successes and failures in cervid conservation, suggest
that there are no universal and global solutions to conservation. The review suggests
that the first priority in conservation must be a detailed understanding of the
autecology of each species. The better the science, the richer the opportunities for
effective intervention, especially where limited means are at hand. In practice this
means that regardless of ongoing conservation efforts, there is a need for continuity
of in basic research, with due regard for geographic species differences in ecology.
Some troublesome foci that need scientific attention are taxonomy, as such has crept
into legislation and can now be subject not to scientific but judicial interpretation.
Predation and cervid security adaptations need attention, as current examples of
predator reintroductions have highlighted deep errors in formerly accepted
assumptions. How to manage cervids is thus not unrelated to how to mange predators.
Here, as in other lines of inquiry the local Pleistocene history of faunas may be
surprisingly relevant. In North America habituation of wildlife is vexing concern.
However, while science is a necessary, it is not a sufficient condition for conservation.
It is necessary to research models of conservation for relevant policies and practices.
Here the nearly century old North American model of Wildlife Conservation offers
important insights. It not only returned wildlife and thus basic bio-diversity to North
America, points to policies essential to conservation as well as the creation of wealth
from wildlife, but also is the only large-scale system of sustainable natural resource
use to date. Using this model as a guide indicates that, globally, the de facto ownership
and use of wildlife by politically potent group of people is the best long-term
protection for wildlife.
(Plenary lecture)
2
Deer responses to global environmental changes.
M. C. Forchhammer
Department of Arctic Environment, National Environmental Research Institute,
Frederiksborgvej 399, POBox 358, DK-4000 Roskilde, Denmark
โIn one sense the conditions of life may be said, not only to cause variability, either
directly or indirectly, but likewise to include natural selection, for the conditions
R 3
determine whether this or that variety shall survive.โ Charles Darwin (On the Origin
of Species, 1859).
Ever since Darwinโs foreseen reflections, we have become increasingly aware of the
importance of abiotic or climatic conditions in forming the evolution of organisms,
their life histories, distributions and population dynamics. Indeed, our awareness
peaked in 1995, when the Intergovernmental Panel for Climate Change summarised
that โthe balance of evidence suggests a discernable human influence on global
climateโ, realising that in addition to direct interactions, we may indirectly, through
our apparent increasing influence on global climate, incur even more severe damage
to the Earthโs ecosystems. Clearly, understanding how climate affects the lives of
organisms is seminal for predicting the effects climate change on ecosystem structure
and functioning. Effects of local weather conditions and their seasonal influence have
been studied intensively in the past. Recently, however, focus has turned to the
potential ecological repercussions following changes in global climate and, during the
last few years, the use of integrative, large-scale climate indices has become somewhat
of an โindustryโ. There are at least two important reasons for this. First, whereas local
weather vary in response to global climate change, large-scale systems, like the North
Atlantic Oscillation, is a major component of global change, which, indeed, may be
an advantage to use when ecological responses have to be interpreted on a large-scale
and, in particular, in a global change context. Secondly, due to its integrative nature,
implicating large-scale climate indices rather than local weather may provide us with
definitive cues of across-species as well as across-taxa responses to global change,
which is central to climate-related issues of biodiversity and conservation. Here, I
selectively review recent studies of deer biology in relation to interannual variations
in the North Atlantic Oscillation (NAO) / Arctic Oscillation. The NAO is a large-scale
seesaw oscillation in atmospheric mass along a meriodinal gradient in the North
Atlantic. By influencing the speed and direction of westerly surface winds across the
Atlantic Ocean, the NAO induces variation in temperature and precipitation in North
America as well as Northern Europe and may, hence, potentially affect the deer
species living in these regions. In particular, I focus on explaining and contrasting the
potential direct and indirect effects of variations in the NAO on phenotypic variation
in life history traits, dynamics of populations and, ultimately, the intertrophic
interactions characteristic of deer species and the environment in which their
embedded.
(Plenary lecture)
4 R
3
Emerging disease in wild and captive Cervids.
M. R. Woodbury and J. R. Campbell
Dept of large Animal Clinical Sciences, Western College of Veterinary Medicine,
University of Saskatchewan, Saskatoon, Saskatchewan, CANADA
Abstract
Global warming and a resurgence of nuclear energy issues are evidence of the
geographical and political pressures that are influencing and altering our world. One
of the consequences of the accelerated changes to the 21st century world is the
emergence and re-emergence of infectious disease in animal and human populations.
Changes to agricultural and wildlife management activities and policies are important
determinants of emerging disease. Furthermore, human activity, or anthropogenic
change, is the single most potent force driving the emergence of these diseases.
Perhaps in response to a global increase in meat consumption, there has been an
increase in the farming of unconventional or non-traditional livestock, including
cervids. Deer farming has created changes in the ecology of deer diseases and has
altered previous concepts of the wildlife/livestock interface. We know very little about
the interactions of wild and farmed cervids or wild cervids and domestic livestock
such as cattle. Likewise we have not examined the importance of cervid product
movements, wild or farmed, in the transmission of disease. We do know that
translocation of wildlife for management purposes and commercial trade in farmed
cervids are human activities that have resulted in pathogen pollution and the spread
of new and old diseases such as chronic wasting disease of cervids and
Mycobacterium avium subspecies paratuberculosis infection, also known as Johneโs
disease. Environmental change and modifications in agricultural methods of food
storage and the artificial feeding of wildlife for hunting and non consumptive purposes
have resulted in the re-emergence in North America of Mycobacterium bovis
infections also known as bovine tuberculosis. This paper examines the emergence of
bovine tuberculosis in elk from Riding Mountain National Park, Canada and in White-
tailed deer in Michigan, USA. The history and epidemiology of chronic wasting
disease in North America is briefly examined as an example of a newly emerging
disease. M. paratuberculosis infections in elk from Point Reyes National Park in
California and Key deer in the National Key Deer Refuge in Florida are discussed as
examples of the importance of understanding disease ecology. One of the answers to
disease emergence is to recognize that there is an ecology of disease in farmed and
wild populations of cervids and for us to carefully consider the consequences of
anthropogenic change to ecosystem health.
Introduction
Global climate changes and a resurgence of nuclear energy issues are evidence of
the geographical and political pressures that are influencing and altering our world.
The consequences of change on such a large scale are more than most of us can
R 5
comprehend but we can understand the manifestations of changes affecting us more
directly such as what we eat and how we produce it. For instance, the world is eating
more meat than ever before. Increased incomes and standards of living in developing
countries are allowing people to eat diets that are higher in protein. World
consumption of livestock products has more than doubled in the past 30 years, driven
mainly by increases in consumption of meat and dairy products in nations such as
China and Brazil, which have accounted for more than half the increase in per capita
meat consumption in developing nations since the 1970's (Holmes, 2001). This shift
in dietary habits has affected the use of agricultural land and changed the way food
animals are produced. There has been an expansion and intensification of livestock
production that has favoured industrial farming systems because they offer an
inexpensive way to produce many animals on a single site, using small amounts of
land close to markets or ports (Holmes, 2001). Even in countries with lots of available
land there has been an increase in โlandlessโ systems using stalls, pens, and feedlots
to intensively raise livestock. The Food and Agriculture Organization of the United
Nations (FAO) has predicted that the proportion of cattle and sheep produced in this
manner in Latin America, sub Saharan Africa, the near East, and North Africa will
more than double by 2030 (FAO, 2000).
The demand for protein and perhaps more importantly the need for a broader
economic base in agriculture in some countries has led to the farming of nontraditional
species of animals such as ostrich, emu, wild boar, crocodiles, guinea pigs and
capybara. Although deer have been farmed in one form or another for millennia, deer
farming is a relatively new agricultural activity to Western countries like Canada, the
United States, New Zealand and Australia. The rise in non-traditional animal
agriculture parallels the burgeoning global trade in bush meat, which is rapidly
becoming a concern to conservationists because of the alarming increase in the
number of threatened species and to world health officials from the equally alarming
risks from zoonotic disease posed by the consumption of bush meat. It has been
estimated that tens of millions of wild animals are shipped each year regionally and
from around the world for food or use in traditional folk medicine (Karesh, 2005). For
instance, live wildlife markets in Guangzhou, China trade in masked palm civets,
ferret badgers, barking deer, wild boars, hedgehogs, foxes, squirrels, bamboo rats,
gerbils, and various snakes and other reptiles as well as domestic dogs, cats and
rabbits (Asia Animals Foundation, 2006).
Intensification of animal agriculture, addition of newly farmed species, and the
trade in wildlife to satisfy the need for protein in human diets are examples of forces
that drive local, and sometimes global, ecosystem changes. One of the consequences
of changes of this nature is the emergence or re-emergence of infectious disease. The
general determinants of emerging infectious disease are described as increased
international travel and trade in animals, changes in human demographics and
behavior, advances in agricultural technology and intensification of the livestock
industry, increased economic development and land use, breakdown of public health
measures and microbial adaptation and change (Daszak et al., 2002). Most of these
determinants are directed by human activities and behaviors and their influence on
ecosystems and the ecology of diseases. Ecology is the study of how organisms
interact with each other and their physical environment. The relationships of wild
cervids with other animals including farmed species and their environment is, to a
6 R
large extent, influenced by humans and therefore the largest part of emerging disease
in cervid populations is driven by anthropogenic changes and activities.
Emerging disease problems can be created or made worse by poor decision making
and planning by wildlife managers and agriculturalists. Robert Hepworth, executive
secretary to the Convention on the Conservation of Migratory Species of Wild
Animals (CMS) said "The proximity of migratory birds to poultry is the outcome of
incorrect planning and faulty development, which have caused the sharing of
important habitats for migratory birds - like wetlands - between wildlife and farms,
with the obvious consequences that we are now experiencing." The sharing of habitats
and the decision to farm new species or to traffic in animals and animal products are
examples of human activities creating new interfaces for disease occurrence. The
wildlife/livestock interface is complicated by the fact that some of the livestock
populations are now also cervids and the emergence of infectious disease in wild
cervids has been made possible, perhaps inevitable, by changes made to the
wildlife/human interface by human activities such as supplemental feeding of wild
deer or deliberate changes to habitat favoring artificially high population densities.
The translocation of cervids and cervid products, either for commercial or wildlife
management purposes, has created enormous opportunities for the spread of disease
between cervid populations. North American examples of emerging cervid disease are
bovine tuberculosis (Mycobacterium bovis) and Johneโs disease (Mycobacterium
paratuberculosis infection) in elk (Cervus elaphus) and white-tailed deer (Odocoileus
virginianus), and the transmissible spongiform encephalopathy of cervids, chronic
wasting disease.
Bovine tuberculosis
Bovine tuberculosis can be found in wild white-tailed deer in a localized area of
the state of Michigan, USA. In 1994, a hunter-killed deer in Michigan was found to
have lesions of tuberculosis, prompting a 1995 survey that isolated M. bovis from
5.1% of the deer sampled (Palmer et al., 2000). Subsequent multi-species sampling
since 1996 has found M. bovis in 19 coyotes, 8 raccoons, 7 black bear, 4 bobcat, 3 red
fox, 2 opossum and 1 domestic cat. Since 2000 there have been 30 positive beef, and
7 positive dairy herds. Also since 2000, there have been 4 positive wild elk found
(Michigan Government, 2006). In 1997, a captive herd of white-tailed deer was found
to be infected and when depopulated, it had a herd prevalence of 12% (Palmer et al.,
2000). Interestingly, the captive herd appears not to have contracted tuberculosis from
interacting with wild deer outside the farm fences, but when the farm was created,
wild deer were purposely trapped inside the perimeter fences as they were erected and
these were used to populate the farm with animals. This was confirmed by DNA
typing of the M bovis organisms found in the farmed deer (Palmer et al., 2000).
Continued surveillance in the local deer population indicate that since intervention
strategies have been employed prevalence rates have fallen to approximately 0.2%
(Michigan Government, 2006).
Presumably the origin of the tuberculosis infection was an interaction between
wild deer and infected cattle. The conventional wisdom of 20 years ago thought that
tuberculosis could not sustain itself in a wild deer population. Prior to 1994 M. bovis
infection had been diagnosed only sporadically in wild North American deer (Rhyan
et al., 1995, Friend et al., 1963, Levine, 1934, Belli, 1962). What happened? Human
R 7
activity in the ecosystem caused the contact rate of infected and uninfected deer to
favor the transmission of disease. It had become common practice in Michigan for
private citizens to supplement the winter food supply of deer on their property with
sugar beets, carrots, corn, apples, and pelleted feed in order to discourage migration
and decrease winter death losses, thereby increasing deer numbers for hunting and non
consumptive tourism. The practice of supplemental feeding increased the population
of deer to focal concentrations of 19 to 23 deer per km
2
(Palmer et al., 2004b). Winter-
feeding not only increased the contact rate among susceptible animals but provided
fomites through which transfer of infective organisms to uninfected deer could occur
(Palmer et al., 2004a). The epidemiological principle at work here involves the basic
reproductive number (R
0
) of disease, which refers to the number of secondary cases
expected from the introduction of one primary case into a completely susceptible
population. R
0
=pcD where p = the probability of infection on contact, c = contact rate
(number of contacts per unit of time), and D = duration of infectiousness (Wobeser,
2006). In order for disease to remain in a population R
0
must be 1. Since the contact
rate is determined primarily by behavior, transmission routes of the pathogen, animal
density, environment etc. in a specific population, it is obvious that anthropogenic
changes in the local ecosystem are responsible for an increased R
0
and the subsequent
maintenance and increase of tuberculosis in wild Michigan white-tailed deer.
Similar circumstances exist in the Greater Riding Mountain Ecosystem of
Manitoba, Canada where wild elk (Cervus elaphus manitobensis) have become
infected with and are maintaining bovine tuberculosis. The difference is that human
activities have not only altered population density and contact rates but have greatly
influenced animal behavior and movement as well. Riding Mountain National Park
(RMNP) is a relatively small reserve (approx. 3000 km
2
) located in a forest-
agricultural transition zone and is almost surrounded by agricultural land. There are
currently about 50,000 cattle in the area around the park with an estimated 2500 elk,
2500 moose (Alces alces), and 5000 white-tailed deer in the greater ecosystem. In the
1950's and 60's bovine tuberculosis was routinely found in cattle that were allowed to
graze inside the park. Even subsequent to a very effective national tuberculosis
eradication program in the 1970's, tuberculosis was occasionally (1981, 1991) found
in cattle herds bordering the park and since 1991 there have been 11 cattle herds
diagnosed with tuberculosis in 4 separate outbreaks (1991, 1997, 2001, 2003) in the
vicinity of RMNP. In 1992 the first naturally tuberculosis infected elk reported in
North America was shot by a hunter in the vicinity of a diseased cattle herd. Between
1998 and 2000, four more infected elk were found in or near the park through a multi-
jurisdictional wildlife surveillance effort that included deer, moose, bear, beaver,
coyote, and raccoon samples. The only non cervid species ever to have been
demonstrated to have Tuberculosis were 2 wolves in the 1960's, discovered
retrospectively in park records of wildlife necropsies. Farmed cervids have not been
affected by tuberculosis in Manitoba. Since 2000, a total of 20 tuberculosis positive
elk and 7 positive white-tailed deer have been discovered within the Greater Riding
Mountain Ecosystem (Copeland 2006, Copeland pers com.). Prevalence is not high
but the existence of the disease is significant because of the impact this tuberculosis
reservoir is having on local cattle production and export trade.
Despite the obvious relationship between cattle, elk, and white-tailed deer disease
near RMNP, researchers know very little about the wildlife/livestock interface in this
8 R
ecosystem and little is known about the ability of elk to transmit M. bovis among
themselves or to other susceptible hosts. In 2002 telemetry collars were placed on elk
from RMNP to examine animal movements and to gather information on the temporal
and spatial aspects of elk interactions with agriculture at the periphery of the park.
Data were also collected to examine environmental and farm management factors that
influence these interactions (Brook et al., 2006). It was widely believed that due to
park management policies elk habitat inside the park has changed and that less
favorable conditions now exist, forcing elk to use agricultural land to feed. Fires, once
commonplace on the prairies and allowed to burn, are now suppressed. Haying,
forestry and cattle grazing activities within the park were stopped years ago. This has
caused areas of the park to be overgrown with forest and shrub cover. Beavers, no
longer trapped in the park, have flooded large areas of elk habitat. It is perceived that
poor quality habitat inside RMNP forces elk out of the park to feed on agricultural
land (Brook et al., 2006). Standing hay and grain crops, and stored hay bales are
regularly consumed and destroyed and fences are damaged. Similar to Michigan,
hunters and outfitters have used food to bait the elk out of the park so that they could
be more easily hunted. Elk accumulate at hay bales left exposed in the fields, at
feeding sites for local cattle, and at bait piles placed by hunters providing opportunity
for transmission of tuberculosis between elk and between elk and cattle through direct
contact as well as feed contamination (Palmer et al., 2004a, 2004b, Brook et al.,
2006).
In both Michigan and in Manitoba the approach to resolving an emerging
tuberculosis problem in cervids has been to modify the conditions that permitted R
0
to increase to disease maintenance levels. This strategy should cause a decrease in the
incidence of new disease and a move towards extinction. In Michigan, the provision
of winter feed supplements has been banned and the number of hunting permits for
deer of both sexes has been increased (Schmitt 2006). In Manitoba, a large public
relations and information campaign aimed at farmers to change livestock feeding and
feed storage practices was undertaken. The result has been that forage harvested in the
area around RMNP is now mostly stored in fenced elk-proof hay yards and there have
been changes to the wildlife baiting and feeding regulations and the level of
enforcement of these. An extension of the cervid hunting season has been made and
there was an increase in the number of hunting permits issued in an effort to reduce
the population of susceptible animals (Brook et al., 2006). In Michigan apparent
prevalence in the core area of the outbreak was 1.2% in 2005, a decrease of 76% since
1995 (Schmitt et al., 2006). Similar results are expected for the RMNP area (Copeland
pers com).
Chronic Wasting Disease
Chronic wasting disease (CWD) was first recognized in 1967 among captive mule
deer (Odocoileus hemionus) that had been assembled at wildlife research facilities in
Colorado from several sources including free ranging populations and a wildlife
research facility in Wyoming (Williams 1980). It wasnโt until 1978 that chronic
wasting disease was identified as a spongiform encephalopathy and at about the same
time the disease was identified in captive mule deer and elk in the Wyoming facility
(Williams et al., 1980, 1982). In the years between 1980 and 1990 CWD was
diagnosed in free ranging elk, mule deer, and white-tailed deer in Colorado and
R 9
Wyoming. Surveillance data and modelling have suggested that CWD may have been
present in free ranging populations for decades before being discovered (Miller et al.,
2000).
The origins and nature of the causal agent of CWD remain contentious issues but
the cause is presumed to be an abnormal prion that perhaps resulted from a
spontaneous change in the configuration of normal prion protein to a resistant form
occurring in a Wyoming mule deer, with subsequent transmission to other deer and
elk in the area. Alternatively, mule deer in the research facility could have been
infected with a cervidadapted strain of scrapie derived from research sheep also
housed at the facility, or even a genetic form of TSE arising in deer, with subsequent
natural transmission (Williams et al., 2002).
In 1996, CWD was discovered on an elk farm in Saskatchewan, Canada and
subsequent disease investigations and tracing of animal movements resulted in finding
several more infected farms. CWD has been found on 40 farms in Saskatchewan and
3 deer farms in Alberta. Approximately 8,000 cervids from infected and trace-out
herds have been killed with 227 testing positive however, there have been no new
cases in Canadian farmed elk in approximately 4 years. Canadian Food Inspection
Agency traceback investigations indicate that CWD was imported into Canada in 1989
through commercial trade in farmed elk from South Dakota (Kahn 2004). CWD was
first diagnosed in the US elk industry in 1997 and since then the disease has been
found in commercial cervid farms in Colorado (1herd), Nebraska (4), Minnesota (2),
South Dakota (8), Montana (1), Kansas (1), Wisconsin (7), Oklahoma (1), and New
York (2) (Chronic Wasting Disease Alliance, 2006). CWD has been found in free
ranging cervids in all of the above states as well as Utah, New Mexico, and Illinois
(Chronic Wasting Disease Alliance, 2006).
Prior to 2000, CWD in free-ranging deer was believed to be limited in the wild to
a relatively small endemic area in northeastern Colorado, southeastern Wyoming and
southwestern Nebraska. However, since 2002, CWD has been found in new areas of
these states as well as those listed above, and including states where there are no
cervid farms. Researchers suggest that the emergence of CWD in farmed and free
ranging cervids are essentially independent events with minimal geographic overlap
and if the epidemics share a common origin, it dates back several decades (Williams
et al., 2002). This is quite possible since, in the early days of elk farming, foundation
stock were often derived from wild populations in what are now known as endemic
areas for CWD.
What sets CWD apart from other TSEs is the fact that it is apparently contagious
and can be spread horizontally from animal to animal or from prion contaminated
environments to cervids in the environment. The incubation period can exceed 36
months, and the clinical course of disease can be protracted (days to months), making
unapparent carriers and prion shedders important to the dissemination of infectious
material in the environment. The determinants of emerging infectious disease
operative in the example of CWD are: increased trade in animals, advances in
agricultural technology, intensification of the livestock industry, increased economic
development and land use, and disease agent adaptation and change. Because of the
Canadian system of individual animal identification, the CFIA were able to trace the
movement of infected animals, and leave no doubt that commercial trade and
translocation of farmed cervids was responsible for the geographic spread of CWD in
10 R
the western Canadian farmed cervid industry. Similar assumptions can be made about
the spread of CWD in the United States cervid industry. What is less clear is the
relationships between disease in farmed cervids and free ranging cervids. There is an
assumption that CWD in the wild has resulted from spillover infection from escaped
animals or interactions between farmed and wild cervids. As far as I know there is
only circumstantial spatial or geographic evidence for this. In fact, very little is known
about the ecology of this disease (on farm or in the wild), the livestock/wildlife
interface, and the interactions between hosts, ecologic and climatic influences, and
societal/transport/commercial factors. We do know that the most common factor
driving the emergence of wildlife infectious disease is the anthropogenic movement
of pathogens into new geographic locations, a phenomenon called pathogen pollution
(Daszak et al., 2001). The translocation of infected hosts either through commercial
trade or wildlife management activities constitutes pathogen pollution. In a broad
sense, so does artificial increases in uninfected host populations through human
activities such that emergence of disease in a new geographic location is supported
(Daszak et al., 2001).
As with bovine tuberculosis in cervids, increased population density resulting from
farming or artificial feeding of wildlife increases the likelihood of direct and indirect
transmission between individuals. In a heavily infected Wyoming research facility,
more than 90% of resident mule deer either died or were euthanatized due to clinical
CWD over a 2year period (Kahn et al., 2004). In the US, CWD was shown to be the
primary cause of adult mortality in at least 2 captive elk herds (71 and 23%) and
prevalences of 59% have been detected at slaughter in elk from CWD affected farms
(Williams et al., 2002).
There is a strong environmental component to CWD ecology in that environmental
contamination is a factor in transmission. Miller et al. (2004) demonstrated that CWD
can be transmitted to susceptible animals indirectly from environments contaminated
by excreta or decomposed carcasses. Mule deer became infected when placed in
paddocks where infected deer carcasses had decomposed in situ almost 2 years earlier
and also in a paddock where infected deer had resided just over 2 years earlier (Miller
et al., 2004). The role of scavengers in the spread of prions in the environment remains
unknown and is an example of the complex possibilities for the role of other
ecosystem factors in emerging diseases like CWD. Similarly, disposal of prion-
infected carcasses resulting from disease eradication efforts on commercial cervid
farms creates environmental concerns. The current Canadian practice has been pit
incineration followed by deep burial but the long-term consequences of this are
unknown.
Johneโs disease
Johneโs disease or paratuberculosis is a chronic intestinal tract disease of ruminants
resulting from Mycobacterium avium subsp. paratuberculosis (MAP) infection. It has
been reported in elk and red deer (Cervus elaphus) as well as white-tailed and mule
deer (genus Odocoileus), axis deer (Axis axis), and fallow deer (Dama dama).
Paratuberculosis organisms have been found in numerous other ungulates, rabbits
(Oryctolagus cuniculus), foxes ( Vulpes vulpes), stoats (Mustela erminea), and ravens
(Corvus corax)(Manning et al., 2003). The organism persists in the environment and
has been recovered from standing water and feces almost a year after contamination
R 11
by infected animals. Disease transmission is easily accomplished via fecal-oral or
transplacental routes and through ingestion of milk or colostrum from infected
females.
In the late 1980's, MAP organisms were isolated from 2 of 5 asymptomatic adult
white-tailed deer killed on a Connecticut cattle farm that had a 6-year history of
bovine paratuberculosis. The authors of the documenting report observed that wildlife
populations had become infected and could therefore serve as continuous sources of
infection to domestic livestock. Their concern was that infected deer migrating to
paratuberculosis-free pastures might seed the soil with organisms that would then
persist in the environment and subsequently infect cattle placed there (Chiodini et al.,
1983).
In 1978, 14 tule elk (Cervus elaphus nannodes) were released at the Point Reyes
National Seashore, California into a fenced area that had been livestock range for
more than a century. In 1979, fecal testing of local dairy herds and previously
introduced axis and fallow deer in the park found that 50% of the dairy herds, as well
as 9.6 % of axis deer and 5.4 % of fallow deer killed in a depopulation effort were
infected with MAP organisms (Riemann et al., 1979). Predictably, 3 tule elk born in
the fenced translocation area showed clinical signs of Johneโs disease in 1981 and
after culling were diagnosed on post mortem with paratuberculosis infection (Jessup
et al., 1981). Since the initial release, the Point Reyes tule elk population has grown
to more than 400 animals and is in excess of environmental carrying capacity. A more
recent report of infection rates in elk from the Park found that MAP organisms were
cultured from 10 of 45 (22%) elk intended for translocation from the infected area to
unfenced areas of the park (Manning et al., 2003). Twenty-two of the 45 elk tested
positive by ELISA or fecal test and were killed leading to isolation of the organism
in 10 animals. Eventually, 18 animals were translocated in spite of the fact that
serology testing for Johneโs is known to be relatively insensitive.
Interestingly, the author cited above reported that the elk tested were all
asymptomatic. Despite the presence of paratuberculosis, the elk population grew
rapidly initially and stabilized over 2 decades with apparently low morbidity and the
infection appears to have had minimal impact on the population. This is confusing
since it is understood that Johneโs disease often profoundly affects captive cervid
herds of many species, often with a much shorter clinical course and in age groups not
usually affected with cattle or sheep herds (Manning et al., 1998).
Clearly, the ecology of Johneโs disease in tule elk at Point Reyes National
Seashore differs from that in other wild populations and commercial cervid farms.
However, we know nothing about the interactions between the high environmental
bacterial burden, a stable host population, possible spillover hosts, and the many other
determinants of Johneโs disease in this ecosystem. It is entirely possible that
equilibrium between Johneโs disease and the tule elk population has been achieved in
this ecosystem such that the host and its immune system have adapted to the presence
of M. paratuberculosis. If a stable Johneโs disease ecology exists it is possibly because
we have allowed it to occur through not interfering. At Point Reyes there is no threat
to the agricultural economy or humans. To date, a causal association between
paratuberculosis and human disease has not been demonstrated. This cannot be said
for tuberculosis caused by M. bovis, and CWD will be always be viewed as a zoonotic
threat because of the association between variant Creutzsfeldt Jakob disease of
12 R
humans and another prion disease of ruminants, bovine spongiform encephalopathy
or mad cow disease.
Another, more enigmatic occurrence of Johneโs disease in Key deer, an
endangered subspecies of white-tailed deer (Odocoileus virginianus clavium) found
only on several islands known as โkeysโ off the southern tip of Florida. Hunting
pressure in the early 1900's reduced the herd to less than 100 deer before legal
protection of the species and the establishment of the National Key Deer Refuge in the
Florida keys increased their numbers to approximately 400 animals by the 1970's
(Quist et al., 2002). However, a 1986 population study suggested that the Key deer
numbers were declining (Humphrey et al., 1986). In 1996 and 1998 single cases of
clinical Johneโs disease were found in deer observed and captured in a residential area
on Big Pine key. A subsequent fecal culture survey for MAP of 96 live-captured deer
yielded one positive deer which was not showing clinical signs at the time of sampling
(Quist et al., 2002). The most intensive culture sampling was done on the small herd
associated with the 2 clinical cases with no positive results. Elsewhere on the key,
only one positive sample was obtained and two subsequent samples from this animal
that remained apparently healthy were negative. These data suggest that large-scale
environmental contamination with MAP is not occurring in the keys. However, a more
or less simultaneous serologic survey of Key deer showed approximately 7% of tested
deer to be possibly infected with MAP, lending support to the hypothesis that MAP
is established, and is perhaps endemic, in Florida Key deer and the ecosystem where
they live.
The origin of the infection in Key deer is unknown. There have been no
commercial livestock operations in the islands since the mid 1900's, but in 1950 a goat
herd existed about 5 km from the location of the index case and local horse owners
sometimes maintain goats as stable companions (Quist et al., 2002). Quist (2002)
suggests that high deer population density, low nutritional quality of native habitat,
and high concentrations of deer at artificial feeding sites might have been contributing
factors to the establishment of MAP infection in this population. What has influenced
the determinants of R
0
such that R
0
1 in this instance is unknown and remains
unstudied using modern epidemiological tools, as does the examination of the role of
anthropogenic change to the ecosystem in the Keys.
Summary
There is little doubt that diseases will continue to emerge in animal populations
including both farmed and wild cervids. In many cases, these emerging diseases are
not necessarily caused by genuinely new pathogens. From the examples described in
this paper, it would seem that many of the emerging diseases in cervid populations are
a direct result of changes in human activities associated with these species. The
intensification of farming of these species, the translocation of species and or their
products resulting in pathogen pollution and human activities which change the
contact rate between both farmed, wild cervids, and other animal populations can
create changes in the ecology of these systems that allow diseases to emerge and
proliferate. Understanding and responding to disease emergence in cervid species
requires a more detailed understanding of the interrelationships between wild and
farmed cervids and other species. There is remarkably little research that helps to
define the contact rates between cervids and farmed animals of a variety of species.
R 13
The study of how cervid species are utilized by humans and their movements of both
animals and products is an important part of understanding this new disease ecology.
Tools such as network analysis and geographical information systems may be
important in examining the role of these factors in disease emergence.
References
Asia Animals Foundation. Species list. (2006) Available from
https://www.animalsasia.org/index.php?module=6&menupos=3&submenupos=5&
lg=en
Belli, L. B. (1962) Bovine tuberculosis in a white-tailed deer (Odocoileus
virginianus). Canadian Veterinary Journal 3:356-358.
Brook, R. K., McLachlan, S. M. (2006). Elk - agriculture interactions in the greater
Riding Mountain ecosystem. Final report to Parks Canada. University of Manitoba,
Winnipeg, Manitoba. Available from
http://www.thegreenpages.ca/tuberculosis/documents/reports/Elk_Agriculture_BR
OOK_FINAL_REPORT.pdf
Chiodini, R. J., van Kruiningen, H. J. (1983) Eastern white-tailed deer as a
reservoir of ruminant paratuberculosis. Journal of the American Veterinary
Medical Association. 182(2):168-169.
Chronic Wasting Disease Alliance. (2006) Learn about CWD, Map updated Feb 8,
2006. Available from http://www.cwd-info.org/index.php/fuseaction/about.map
Copeland, S. (2006) Timeline of bovine tuberculosis (TUBERCULOSIS) in
Canadian and Manitoba cattle and bison. Manitoba Agriculture, Food and Rural
Initiatives, Winnipeg, Manitoba. Available from
http://www.gov.mb.ca/agriculture/livestock/anhealth/jaa04s02.html
Daszak, P., Cunningham, A. A., Hyatt, A. D. (2001) Environmental change and the
emergence of infectious diseases in wildlife. Acta Tropica 78:103-116.
Daszak, P., Cunningham, A. A. (2002) Emerging infectious diseases; a key role for
conservation medicine. Pages 40-61 in A. A. Aguirre, R. S. Ostfeld, G. M. Tabor,
C. House, and M. C. Pearl, editors. Conservation medicine: ecological health in
practice. Oxford University Press, New York.
Food and Agriculture Organization of the United Nations (FAO), Global
Perspectives Unit. (2000) Agriculture: Towards 2015/30. Technical Interim Report.
Rome: FAO.
Friend, M., Kroll, E.T., Gruft, H. (1963) Tuberculosis in a wild white-tailed deer.
New York Fish and Game Journal 10:118-123.
14 R
Holmes, K. (2001) Carnivorous cravings: charting the worldโs protein shift. In:
Earth Trends World Resources Institute. Available from
http://earthtrends.wri.org/features/view_feature.php?theme=8&fid=24
Humphrey, S.R., Bell, B. (1986) The Key deer population is declining. Wildlife
Society Bulletin 14:261-265.
Inch, C. (2003) Chronic wasting disease (CWD) in Canada - Current situation in
Canada. Canadian Animal Health Net Bulletin 8:28-29.
Jessup, D.A., Abbas, B., Behymer, D. (1981) Paratuberculosis in tule elk in
California. Journal of the American Veterinary Medical Association.
179(11):12521254.
Kahn, S., Dube, C., Bates, L., Balachandran A. (2004) Chronic wasting disease in
Canada: Part 1. Canadian Veterinary Journal. 45(5):397404.
Karesh, W. B., Cook, R. A., Bennett, E. L., Newcomb, J. (2005) Wildlife trade and
global disease emergence. Emerging Infectious Disease Online. Available from
http://www.cdc.gov/ncidod/EID/vol11no07/050194.htm
Levine PP. (1934) A report on tuberculosis in wild deer. Cornell Vet 24:264-266.
Manning, E. J., Steinberg, H., Rossow, K., Ruth, G. R., Collins, M. T. (1998)
Epizootic of paratuberculosis in farmed elk. Journal of the American Veterinary
Medical Association. 213(9):13-22.
Manning, E. J., Kucera, T. E., Gates, N. B., Woods, L. M,, FallonMcKnight, M.
(2003) Testing for Mycobacterium avium subsp. paratuberculosis infection in
asymptomatic freeranging tule elk from an infected herd. Journal of Wildlife
Diseases. 39(2):323328.
Michigan Government (2006) Emerging disease issues. Diseases that may affect
humans or animals. Wildlife surveillance table, March 6 2006. Available from:
http://www.michigan.gov/documents/WildlifeTUBERCULOSISSurveillanceSumm
ary_119911_7.pdf
Miller, M. W., Williams, E. S., McCarty, C. W., Spraker, T. R., Kreeger, T.J.,
Larsen, C. T., Thorne, E. T. (2000) Epizootiology of chronic wasting disease in
freeranging cervids in Colorado and Wyoming. Journal of Wildlife Diseases.
36(4):67-69.
Miller, M. W., Williams, E. S., Hobbs, N. T., Wolfe, L. L. (2004) Environmental
sources of prion transmission in mule deer. Emerging Infectious Disease
10(6);1003-1006.
R 15
Schmitt, S. M. (2006) Management of bovine tuberculosis in Michigan deer.
Michigan government emerging disease issues. Diseases that may affect humans or
animals. Available from:
http://michigan.gov/emergingdiseases/0,1607,718625804_2581175930,00.html
Palmer, M. V., Whipple, D. L., Payeur, J. B., Alt, D. P., Esch, K. J., BruningFann,
C. S., Kaneene, J. B. (2000) Naturally occurring tuberculosis in whitetailed deer.
Journal of the American Veterinary Medical Association. 216(12):19211924.
Palmer, M. V., Waters, W. R., Whipple, D. L. (2004a) Shared feed as a means of
deertodeer transmission of Mycobacterium bovis. Journal of Wildlife Diseases.
40(1):8791.
Palmer, M. V., Waters, W. R., Whipple, D. L. (2004b). Investigation of the
transmission of Mycobacterium bovis from deer to cattle through indirect contact.
American Journal of Veterinary Research 65(11):14831489.
Rhyan, J.C., Aune, K., Hood, B., Clarke, R., Payeur, J., Jarnagin, J., Stackhouse, L.
(1995) Bovine tuberculosis in a free-ranging mule deer (Odocoileus hemionus)
from Montana. Journal of Wildlife Diseases 31:432-435.
Riemann, H., Zaman, M. R., Ruppanner, R., Aalund, O., Jorgensen, J. B.,
Worsaae, H., Behymer, D. (1979) Paratuberculosis in cattle and freeliving exotic
deer. Journal of the American Veterinary Medical Association. 174(8):841843.
Williams, E. S., Miller, M. W. (2002) Chronic wasting disease in deer and elk in
North America. Revue Scientifique et Technique. Off Int Epiz. 21(2):3051.
Williams, E. S., Young, S. (1980) Chronic wasting disease of captive mule deer: a
spongiform encephalopathy. Journal of Wildlife Diseases. 16: 89-98.
Williams, E. S., Young, S. (1982) Spongiform encephalopathy in Rocky Mountain
elk. Journal of Wildlife Diseases. 18: 465-71.
Wobeser, G. A. (2006) Transmission and perpetuation of infectious disease. Pages
105-123 in Essentials of disease in wild animals. Blackwell Publishing, Ames,
Iowa.
(Plenary lecture)
16 R
4
Seasonal versus non-seasonal reproduction in deer: From the arctic
to the tropics.
G. A. Bubenik
Department of Integrative Biology, University of Guelph, Guelph, Ontario, N1G 2W1,
Canada
Deer are found in all geographical zones, from the tropic climate to the high arctic.
The richest variety of cervid species are found in Southeast Asia and South America.
The most adaptable deer are white-tailed deer, which are distributed from Northern
Canada to Southern Peru. Smaller species with shorter gestation periods, inhabit
mostly tropical and subtropical regions. Bigger cervids, exhibiting longer gestation
periods, generally live in the temperate and boreal regions. Deer ancestors, which
developed in the relatively warm Miocene period in both Europe and Asia, were most
probably aseasonal breeders. The changing climate in the Pliocene and Pleistocene
and the migration of cervids to temperate and boreal regions forced them to become
a short-day seasonal breeders. However some relics of that ancestral reproductive
pattern still persist. Whereas most tropical and subtropical species exhibit
asynchronous reproductive cycles independent of the photoperiod, most temperate and
boreal cervids exhibit annual rutting season, synchronized by light. Most temperate
species exhibit seasonal breeding characterized by a concise rut and parturition
periods. The activation of seasonal reproduction begins as a response to changes of
environmental photoperiod and continues as a cascade of hormones which include
melatonin, prolactin, luteinizing hormone (LH), follicle stimulating hormone (FSH)
and testosterone. Cervids are capable to respond to photoperiodic clues if the
difference between summer and winter solstice is at least one to one and half hours.
That global transition zone (separating tropical from temperate cervids) corresponds
to latitudes of 14-18
o
N or S. A substantial evidence indicates that ancestors of
temperate cervids might have had two rutting periods per year. Two peaks of
testosterone in blood (smaller in the spring, larger in the fall) were found in plasma of
several boreal cervids species. In addition Southern pudu (Pudu puda), living in
central Chile, exhibits only one rut but two equally high peaks of testosterone and LH,
whereas his Northern cousin (Pudu mephistophiles), living in the tropical regions of
Peru, have two rutting seasons per year. Similarly, two peaks of reproductive
hormones (one corresponding to antler mineralization, the other to the rut) are present
in the male European roe deer. Many deer males (irregardless of their geographical
distribution) are fertile for a better part of the year and are capable of mating, if the
females are in the heat. Some tropical and subtropical cervids are year round breeders,
others are seasonal but non-photoperiod breeders. Their rutting and parturition periods
are determined by environmental factors other than light, such as the onset of the rainy
season. Temperate deer species, translocated to the Southern hemisphere, synchronize
quickly with the local photoperiod. However, in some translocated tropical and
subtropical species their reproductive and antler cycles became either asynchronous
R 17
(exhibiting year round breeding) or they maintain their original seasonal pattern which
is unsuitable for the survival of their offspring.
(Plenary lecture)
5
Fallow deer, lekking and alternative mating strategies in San
Rossore, Italy: insights from a long term study.
M. Apollonio
Department of Zoology and Evolutionary Genetics - University of Sassari, Italy
Throughout the analysis of long-time series census data (1984-2003) and by means of
radiotelemetry (1997-2001; 23 females and 25 bucks), we assessed fallow deer spatial
and habitat sexual segregation on different spatial and temporal scales in the San
Rossore Estate (Italy). The combination of different studies performed in the same
area allows us to show that a detailed analysis can outline a situation in which more
than one theory is valid for explaining sexual segregation in the same species, as
probably several factors act on different temporal and spatial scales. From 1984 to
2003 females gradually abandoned the eastern disturbed sector of the Estate, the only
area affected by human presence (increased year by year from 1984), to the detriment
of the western undisturbed sector. Males remained in the eastern sector, in spite of the
increasing disturbance, supporting the predation risk hypothesis (large spatial and
large temporal scale). From 1989, males gradually increased their presence in the
disturbed sector, as they benefited from lower female density, avoiding areas with
higher female density, supporting therefore the indirect competition hypothesis (large
spatial and large temporal scale). Intersexual competition was pronounced in a small
area inside the undisturbed sector affected by habitat modification during the 1980s,
when a scrub area was converted to an open pasture, and this phenomenon further
incited males to leave this area and reach the disturbed sector, supporting again the
indirect competition hypothesis (small spatial and large temporal scale). As a final
result of this long term process, during the last five years we showed that males used
consistently disturbed areas, both during the day and the night; instead, females
frequented eastern disturbed areas only during the night, when human presence was
absent. Therefore, large scale segregation recorded during the day disappeared during
the night, when females usually reached the eastern sector, supporting again the
predation risk hypothesis (large spatial and small temporal scale). Nevertheless, sexes
segregated on a small scale inside this area, as they showed different habitat choices,
supporting the forage selection hypothesis (small spatial and small temporal scale).
(Plenary lecture)
18 R
6
Recent progress in antler regeneration and stem cell research.
C. Li and J. M. Suttie
AgResearch New Zealand Ltd, Invermay Agricultural Centre, Mosgiel 9007, New
Zealand
Epimorphic regeneration is the โHoly Grailโ of regenerative medicine. Research
which aims to investigate different models of epimorphic regeneration is essential if
a fundamental understanding of the factors underpinning this process are to be
established. Deer antlers are the only mammalian appendages that are subject to
annual epimorphic regeneration. Therefore, antlers offer a unique opportunity to
explore how nature has solved the problem of mammalian organ regeneration.
However, establishment of an antler model for epimorphic regeneration study in the
past has been painfully slow. With the advancement of the modern technologies, antler
biologists in recent years have made significant progress in this field. Through detailed
morphological and histological examinations, we advanced a hypothesis that antler
regeneration may not be a blastema-based process as previously thought, but rather
a stem-cell based process. Subsequently, we set up experiments to test this hypothesis.
Using IHC and in situ hybridisation techniques, we provided evidence that the mode
of antler regeneration is not compatible with blastema-based epimorphic regeneration.
Through tissue deletion and membrane insertion, we precisely located the tissue type
from which antler regeneration takes place, i.e. pedicle periosteum. Pedicle periosteal
cells express crucial embryonic stem cell markers, for example CD9, Nanog,
telomerase and a low level of Oct4. In addition they have the potential to differentiate
into multiple cell types, such as chondrocytes, adipocytes and possible neural cells,
convincingly demonstrated that they meet the criteria for adult stem cells. Therefore,
we conclude that antler regeneration is not a blastema-based, but rather a stem cell-
based epimorphic process. Antler stem cells exclusively reside in pedicle periosteum.
In order to study the regulation of antler regeneration, we constructed a subtracted
cDNA library using tissues from the antler growth centre. From this library, we
identified 331 ESTs similar to those with no known function and 17 novel genes.
Whether antler regeneration is realised through recapitulation of developmental
processes, unique antler-specific factors or novel regulation system remain to be
determined in the future research.
(Plenary lecture)
R 19
7
Conservation of tropical deer: what does the future hold?
W. J. McShea
Smithsonian Institution, CRC, 1500 Remount Rd., Front Royal, VA 22630, USA
There are 4 stages of conservation: exploration, science, policy/education, and
management. Whereas, temperate deer populations are often in a management mode,
we still lack basic scientific information for most tropical deer species. A recent
volume of deer biology contained 487 references for temperate species and 69
references for tropical species, 36 of which concerned captive populations. Whereas,
deer management in developed countries has generated significant funds to maintain
and enhance most deer populations, the stark needs for tropical species are unmet by
international organizations. To be sustainable, deer conservation in the tropics must
move to the management level, yet we still lack basic knowledge to make the
transition. Tropical deer conservation currently focuses on 2 factors; suitable habitat
and adequate protection. Suitable habitat is assumed for most deer species, as they are
generalist browsers or grazers. Within Southeast Asia, however, increased pressure
for agricultural production (i.e. rice, sugar cane, and palm oil) has removed significant
amounts of lowland forests. Habitat is not the sole limiting factor, as a recent analysis
for Cervus eldi reveals most suitable habitat is unoccupied. Few governments provide
sufficient protection of low density deer populations. At this time, international aid
would best be focused on supplementing patrolling staff salaries. Training of reserve
staff should emphasize that effective conservation is proactive management of all
resources available. Use of fire and water as habitat management tools should follow
the successful models for bovid management in Southern Africa. Scientists and
managers must be willing to engage in the third stage of conservation (i.e. policy and
education) or there is little hope for moving to a sustainable, management stage for
tropical deer species.
(Plenary lecture)
20 R
Deer management
22 R
8
Three years of roe deer (Capreolus capreolus) radio-tracking in a
Mediterranean environment.
A. J. Ferreira and C. Silva
Instituto Ambiente e Vida - Universidade de Coimbra. Largo Marques do Pombal.
3004 -517 Coimbra. Portugal.
During three years, several roe deer were equipped with radio collars and followed in
summer and winter periods. On those tracking periods several fixes were obtained
(about 30 fixes/animal/season) and the home ranges calculated using MCP and Kernel
models. This study was made in a mountain area (Serra da Gardunha), localized in the
centre of Portugal, in a Mediterranean environment with hot and dry summers and
mild winters. Annual. seasonal home-ranges and it evolution were compared with
results obtained in other populations in different habitats condition. When consider the
distribution of the resources in the study area we can demonstrate that the roe deer
present a spatial behaviour, different from the central and north populations and
adapted to the Mediterranean climate constrains.
(Oral presentation.)
9
Seasonal home range shift of red deer in a forest-agriculture area,
Hungary.
L. Szemethy, Zs. Birรณ, K. Katona, K. Mรกtrai, Sz. Orosz, and N. Bleier
St. Istvรกn University, Department of Wildlife Biology and Management, Pรกter Kรกroly
u. 1., Gรถdรถllล 2103, Hungary
A long-term radiotelemetry study on red deer (Cervus elaphus Linnรฉ, 1758) has been
carried out in a lowland forest-agriculture area in Hungary between 1993-2006.
Previous observations suggested seasonal changes in population distribution between
forested and agricultural habitat. Red deer concentrated in the forest during winter, but
they appeared in the agricultural field during the vegetational period. We investigated
the ranging behaviour behind this phenomenon. To do so two alternative hypotheses
were made: home range expansion and home range shift. Weekly radiotelemetry
localisations revealed that nine of 28 hinds showed a clear home range shift from the
forest to the agricultural area for a prolonged time during the vegetational period. The
remaining portion of the animals used a home range within the forest throughout the
entire year. Diet composition analysis using indicator plant species showed that
neither daily passages between habitats, nor home range expansion exists. These
R 23
ranging behaviours were stable year on end, hence, if an animal shifted one year it
shifted again in consecutive years and vice versa. We suggest, that there is no feeding
reason for seasonal home range shift. In another study area, where radiotelemetry
study on red deer was carried out between 2000 and 2006, we revealed daily passages
between forested and neighbouring agricultural areas. Use of agricultural fields by red
deer is influenced by the scale of the patchiness of landscape including forested and
agricultural areas. Our results could be useful for the successful management of red
deer populations in such complex habitats and to decrease agricultural damage.
(Oral presentation.)
10
Space use patterns of Persian fallow deer following reintroduction.
S. Bar-David
1
, D. Saltz
2
, A. Dolev
3
, A. Perelberg
4
, and T. Dayan
5
1
The Institute of Evolution, Haifa University, Israel
2
Department of Desert Ecology, BIDR, Ben Gurion University, Israel
3
The Mammal center, The Israel Society for Protection of Nature, Israel
4
Department of Evolutionary and Environmental Biology, Haifa University, Israel
5
Department of Zoology, Tel Aviv University, Israel
The ability to disperse and populate new areas is an essential condition for long-term
survival of reintroduced populations. Consequently, understanding the spatial
dynamics of reintroduced populations is crucial. We studied the movement patterns
of 70 radio-collared Persian fallow deer (Dama mesopotamica) (53 f, 17 m), of 120
deer reintroduced to Israel. Deer were released from the same habituation enclosure,
in bi-annual releases, on 10 occasions, starting 1996. We obtained 2-3 radio locations/
/individual/week for up to five years and characterized annual home ranges using GIS.
The spatial dynamics were characterized by: (1) Short distance movements (up to 1.5
km from the release site) until home range establishment, mainly of deer from early
releases; (2) long distance movements, up to 15 km until home range establishment,
mainly of deer from latter releases; and (3) spatial shifts in annual home ranges away
from the release site, which occurred usually within the first three years post release.
The radial expansion of the population during the first 5 years of the project was 0.5-
1.5 km/year, slower than that of other deer species. While population growth over time
was linear (mostly due to repeated releases), the area occupied by deer increased
exponentially, from 6 km during the first year to 47 km during the fifth. This spatial
expansion was largely due to long distance movements from the release site to new
unpopulated areas, which were frequent during the last years. Radiation of
reintroduced Persian fallow deer is driven mostly by animals from later releases. One
possible benefit of multiple releases is increased rates of population expansion.
(Oral presentation.)
24 R
11
Population size and demographic variables of red deer in Bydgoszcz
National Forest, central Poland.
P. Beszterda
Regional National Forest Headquarter in Toruรฑ, Mickiewicza 9, 87-100 Toruล,
Poland
During February 2006, population size of red deer (Cervus elaphus L.) inhabiting
Bydgoszcz National Forest (76.8 thousand ha) was estimated. In 5 Forest Districts
(Bydgoszcz, Cierpiszewo, Gniewkowo, Solec Kujawski and Szubin) including 25
hunting districts, 120 line transects (420 km total length) were established. During 5
consecutive days, the number of daily snow tracks were recorded. Using relationship
between daily snow track density index (T/km) and absolute population density
(N/1000 ha), population density and numbers for every hunting district were
calculated. The everage number of red deer estimated from the 5 days tracking period
was equal to 3850 animals i.e., 50.1 individuals/1000 ha of forest. In September 2005,
data on population sex ratio and the autumn recruitment rate were collected by direct
observation of 959 animals. Sex ratio i.e. females per one male was equal 1: 1.50. The
autumn recruitment rate amounted to 48.2 calves per 100 females. It was concluded
that the official hunting statistic data based upon so called round year observation,
underestimated 3 times number of red deer calculated in presented work.
(Oral presentation.)
12
Management of red deer in Poland: field data versus official
hunting statistic.
B. Bobek
1
, T. Mamok
2
, J. Mikoล
3
, W. Rembacz
4
, A. Standio
5
, and R. Wasilewski
6
1
Department of Ecology, Wildlife Research and Ecoturism, Pedagogical University
of Cracov, Podbrzezie 3, 31-054 Krakow, Poland
2
2
Rudziniec Forest District, Leลna 4, 44-160 Rudziniec, Poland
3
Wejherowo Forest District, Sobieskiego 247 B, 84-200 Wejherowo, Poland
4
Myลlibรณrz Forest District, Dworcowa 2, 74-300 Mysliborz, Poland
5
Zdrojowa Gรณra Forest District, Poznanska 126, 64-920 Pila, Poland
6
Regional Directorate of State Forest in Gdaลsk, Rogaczewskiego 9, 80-804 Gdaลsk,
Poland
Management of red deer in Poland is based upon population census including
estimating number of the animals in hunting districts, sex ratio of adults, age structure
of stags and annual recruitment rate. We have compared population numbers and
R 25
structure based upon official hunting statistic data with data collected during 1998-
2006 by line/strip transect technique and by standard observations. Analysis of 155
Forest Districts (FD) of the total area of 2.48 mln ha, showed that only in 14 FD
hunters data were ยฑ15% different than data collected by line/strip transect method. In
98 FD number of red deer was higher than official hunting statistic data and this error
ranged between 15-50%. Much higher errors i.e., 50-100% and over 100% were found
in 31 FD and in 12 FD respectively. Data from direct observation cards showed that
sex ratio of adults (females/males) was calculated as 1:1.68, while according to
hunters this parameter was equal to 1:1.33. Hunters report there are 9.7% stags over
10 years old, but field data indicated only 4.6% such age class in the population of
males. The described situation is result of the present Hunting Low that allows hunters
to use quess-estimate data for red deer management in Poland. Such non-professional
management resulted in overabundance of deer in many regions of the country and
drastic decline of high trophy stags in the population of red deer. The management of
red deer has to be based upon data collected by hunters in field using objective and
reliable methods. This basic data should be processing in Forest Districts by
professional service and the hunting clubs would obtain back population census and
harvest plan. However the presented here feed-back mechanisms of data flow can be
introduced if the present Polish Hunting Low will be changed.
(Oral presentation.)
13
Over-abundance of deer: Is shooting the answer?
D. C. MacMillan
University of Aberdeen, Scotland
Shooting is an important population control measure. However, in many regions
demand for hunting is declining and it is not clear if shooting can control expanding
deer populations. The issue is particularly acute in the Highlands of Scotland, where
over 2 million hectares of land are owned as sporting estates for shooting deer but
where the population of red deer has more than doubled over the last 30 years,
threatening native woodlands and other protected nature conservation sites. The Deer
Commission for Scotland has identified voluntary shooting effort as the preferred
approach to the problem but in 2003-2004 less than one-third of all local cull targets
were met and culling effort on private land has declined. The aim of this paper are to:
i) investigate the constraints on deer shooting from a landowners perspective and ii)
suggest future strategies for encouraging landowners to shoot more deer. A statistical
analysis of data derived from interviews and a mail survey of landowners is used to
understand the role of motivation and objectives, attitudes to nature conservation and
the regulatory system, as well as resource constraints such as labour on culling
activities. Key findings are: 1) traditional deer management objectives seek to satisfy
demand for trophy stags by maintaining high population densities and this over-rides
26 R
other objectives; 2) financial rewards from increasing culling rates are meagre; 3)
shooting efficiency of some estate employees is low and may reflect poor motivation
and 4) there is no direct public funding to assist private estates with culling or with the
costs associated with developing markets for venison or commercial shooting. It is
suggested that a possible future strategy based upon better support for marketing of
venison and shooting opportunities, together with better incentives and training for
estate employees might be partially effective. However. resistance to greater culling
is likely to among estates where traditional values persist and where employees lack
incentives to intensify culling operations.
(Oral presentation.)
14
Slaughter records as a body condition indicator or reindeer - How
can records be improved?
A. Olofsson, B. ร
hman, and ร. Danell
Reindeer Husbandry Unit, Swedish University of Agricultural Sciences, Uppsala,
Sweden
Reindeer and caribou (Rangifer tarandus ssp) population densities are known to
fluctuate in cyclic patterns. Pasture quality is one important factor that determines
these fluctuations (Klein, 1968, Reimers, 1997). In semi-domesticated reindeer herds,
the fluctuations are usually not as dramatic as in many wild populations, but they still
are large enough and cause problems and an unstable economic situation for the
reindeer herder. Knowledge of changes in pasture quality is essential in order to not,
because of management inabilities, make management decisions that drive large
fluctuations. Indicators of pasture quality that can be monitored continuously are
therefore important.
During the snow-free season reindeer herding is usually extensive, using wide
ranges. Direct monitoring of pasture quality on these ranges would be expensive and
time consuming, if possible at all. Since the body condition of reindeer in early winter
is affected mainly by pasture quality during the preceding summer (Adamczewski et
al., 1987, Skjennerberg and Slagsvold, 1968), long-term changes of the general body
condition in the herd should reflect the long-term changes of the pasture. The
condition of the reindeer in the early winter could therefore be a good indirect
indicator of the condition of the ranges used during the snow-free season. The main
slaughter period for reindeer is usually in the early and midwinter (November to
January) and information on carcass size and quality might thus be used to assess the
condition of the reindeer at this time.
The objective of this study was to investigate the possibilities to of using and
improve information from carcass records as an indicator of animal condition.
Records from 696 reindeer slaughtered in the winter 2002/2003 were included in
the study. The records received from the slaughterhouses included herding district,
R 27
carcass weight, fatness and conformation (classified according to the EUROP system)
and animal category (calf, female and male adult). In addition three body size
measurements (back-, radius- and jaw length), sex of calves and age class of adults
(yearlings or older adults) were recorded.
The fatness and conformation classification was transformed into normal
distribution using normal score transformation. Principal component analyses (PCA)
where used for assessing the relationships among weight, the body size measurements,
fatness score and conformation score without needing to define dependent and
independent variables. General linear models (GLM) were used to investigate the
possibilities to use improve precision of body weight, by adjusting for body size, as
a body condition indicator and to investigate the gain in precision of differing between
sex and age categories.
The PCA showed that the relationships among the variables differed between age
and sex categories. The linear models showed similar results since the size and
significance of the fixed effects on weight differed between categories. Male and
female calves were most alike and differed from the older animals. There was a clear
difference between male and female yearlings, where male yearlings showed more
similarities to calves than female yearlings, which more resembled female adults. Yet,
discriminating between female yearlings and older females improved precision of
adult carcass weight records by reducing standard deviation (SD) with 15,5 %. The
results also showed that discriminating between sexes for calves reduced SD of calf
carcass weight records with 36,2 %. It is hence essential to differ between females and
males when calf records are used as body condition indicators, since varying
proportions of the sexes will easily cause a bias.
All body size measurements were strongly correlated within animal age and sex
category (r = 0.78 โ 0.88) and all measurements gave almost similar results when
adjusting weight for body size. Adjusting weight for the different body size
measurements reduced SD of weight records with 8 โ 30 % (exemplified with
adjustment for back length in Tab. 1).
Table 1. Reduction of standard deviation for weight by adjustment for back length
N
Mean
carcass
weight (kg)
SD before
adjustment
(kg)
Remaining
proportion of SD
after adjustment
Female
Calf
103 20,5 2,7 72 %
Male Calf 312 22,7 2,8 71 %
Female
Yearling
44 28,1 3,5 91 %
Male
Yearling
77 30,7 3,0 79 %
28 R
N
Mean
carcass
weight (kg)
SD before
adjustment
(kg)
Remaining
proportion of SD
after adjustment
Female
Adult
159 33,6 3,7 89 %
High correlations between the body size measurements and weights of calves
suggest that weight gain and skeleton development could be considered as parallel
indicators of animal body condition affected by range conditions during the last
season. Therefore, adjusting calf carcass weight for body size would likely diminish
its power as indicator of range conditions. Adult body size, in contrast, is affected by
growth conditions during several previous seasons. Hence adjusting carcass weight
of adults for body size can prevent confusion of range conditions during several
seasons. Instead skeleton size per se may reflect range conditions over a longer time
period.
References
Klein, D. (1968) The introduction, increase, and crash of reindeer on St. Matthew
island. Journal of Wildlife Management 32(2):350-367.
Reimers , E. (1997) Rangifer population ecology: a Scandinavian perspective.
Rangifer 17 (3):105-118.
Adamczewski, J. Z., Gates C. C., Hudson, R. J. and Price, M. A. (1987) Seasonal
changes in body composition of mature female caribou and calves (Rangifer tarandus
groenlandicus) on an arctic island with limited winter resources. Canadian Journal of
Zoology 65(5):1149-1157.
Skjennerberg, S. and Slagsvold, L. (1968) Rein driften og dens naturlag.
Universitetsforlaget, Oslo.
(Oral presentation.)
R 29
15
The management of reindeer in the Mongolian Tsaatan culture.
J. C. Haigh and M. G. Keay
Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon, SK
S7N 5B4, Canada
The Itgel Foundation, 1243 Arroyo Chico Dr, Boulder, CO 80302, USA
The Tsaatan (or Dukha) peoples of north-western Mongolia are one of the few
remaining reindeer herding cultural groups in the world. They are a migratory people
who live and move with their reindeer in the Eastern Sayan mountains that span the
Siberian and Mongolian border. The use of reindeer as transport animals is a
dominating feature of the Tsaatanโs tradition of reindeer husbandry, as is milking deer
for the purpose of making various dairy products consumed by Tsaatan individuals.
The spiritual significance of reindeer in the Tsaatanโs shamanistic traditions plays an
important role in pastoral activities and trends. The close association between the
people and their stock has led to a marked degree of domestication, and the animals
seldom stray far from camp. It is common for animals to be released from their tethers
at dawn, move off to feed for a few hours, and then return to camp without coercion
by about mid-day. Others are herded back in late afternoon in the company of
community members who ride out on saddled reindeer to round them up. During field
trips throughout three years (2002, 2004, 2005), we conducted interviews with herders
in the two distinct areas where families of this community reside. Like most of the
traditional reindeer-herding peoples of the world, the Tsaatan face many challenges
as they struggle to maintain their ancestral lifestyle. One example is the erosion of the
hunting rights that the Tsaatan have enjoyed for centuries, possibly millennia.
Traditionally the Tsaatan have not used reindeer as a primary source of meat, relying
instead on several wild game species such as moose, red deer, wapiti, roe deer, bear,
lynx, wild boar and others. But recent shifts in socio-economic systems stimulated by
Mongoliaโs transition from socialism to democracy in 1991 has led to an increase in
hunting pressure from non-subsistence hunters outside the community, contributing
to the reduction of wildlife populations, and tightened government control over
hunting. On top of this, wolf predation on the herds, especially of the young calves,
has contributed to a decline in reindeer numbers.
(Oral presentation.)
30 R
16
Can supplementary feeding improve productivity in reindeer
husbandry?
B. ร
hman and ร. Danell
Swedish University of Agricultural Sciences, Reindeer Husbandry Unit, Uppsala,
Sweden
Reindeer (Rangifer tarandus tarandus) in Sweden are semi-domesticated and
managed by their owners. They normally graze on natural pastures in forest and
mountain areas all year. Artificial or supplementary feeding is only used occasionally,
mainly during gathering and migration or when hard snow or ice crust makes the
ground vegetation unavailable. In some areas, feeding is used as a measure to reduce
radioactivity prior to slaughter. Reindeer are only rarely fed with the main purpose to
improve their body weights before slaughter.
Artificial feeds for reindeer are mainly grain-based pellets, hay or grass silage or
a combination of these. Hay or silage has proven to be unsuitable as the only feed to
reindeer for longer periods (Nilsson et al., 2000). The costs for feeding reindeer are
high, and the weight gain of the animals is usually low. Reindeer calves fed pellets
during winter will normally not gain more than 100-150 gram live weight
(corresponding to 50-75 gram carcass weight) per day, if fed ad libitum, while eating
about 1,5 kg dry feed (15 MJ) per day (Jacobsen et al., 1977; ร
hman, 1996; Nilsson
et al., 2000 and own recent results). The cost for feed to reindeer calves fed ad libitum
will be about 0.3 euro per animal and day. With the current Swedish prices on reindeer
meat (about 5 euro per kg carcass weight), the animals need to gain at least 120 g live
weight per day, on average, to compensate for the costs for feed. Costs for labour,
transport, equipment etc. will not compensated for if the weight gain (or the meat
price) is not considerably higher.
During harsh conditions, feeding could be the only way to prevent reindeer from
starving and eventually dying. Feeding during critical periods, when grazing
conditions are unfavourable, may also improve the general condition of the reindeer
herd. The benefit on production depend on the general condition of the herd as well
as the severity of the grazing conditions, and is thus difficult to assess. However, if
there is a risk of animals starving and dying the animal welfare aspect has also to be
taken into consideration.
The average condition of adult females has proven to be fundamental for calf
production and survival (Rรถnnegรฅrd et al., 2002). Finnish studies have shown that the
conditions of lichen pastures in winter increases the number of calves per adult female
the following winter (Kojola et al., 1995) and that supplementary feeding with hay
improves the body weights of both calves and females the following autumn (Helle
& Kojola, 1994). Supplementary feeding during winter has also been shown to
improve milk production and calf body mass (Jacobsen et al., 1981). Improving the
condition of females by feeding during winter may thus be an effective way to
improve the production of the herd.
We have simulated the effect of feeding on reindeer body weight during varying
R 31
grazing conditions and at different times of the year to assess the possible long-term
effect on production. In herds with initially low production, the productivity could be
significantly improved by feeding females during late winter and spring. There is little
economic reason to feed reindeer prior to slaughter in order to improve their body
weight and carcass quality. In addition, it has been shown that feeding changes the
chemical composition (Wiklund et al., 2001) and taste (Wiklund et al., 2003) of
reindeer meat, which could be negative. There is also a risk that extensive use of
feeding in reindeer management will change the image of reindeer meat as natural
product and thus be negative for marketing. We therefore believe that artificial feeding
of reindeer should be made only in special situations where it has a proven positive
effect for the production.
References
ร
hman, B. (1996) Effect of bentonite and ammonium-ferric(III)-
hexacyanoferrate(II) (AFCF) on uptake and elimination of radiocaesium in
reindeer. Journal of Environmental Radioactivity 31:29-50.
Helle, T., Kojola, I. (1994) Body mass variation in semidomesticated reindeer.
Canadian Journal of Zoology 72:681-688.
Jacobsen, E., Bjarghov, R. S., Skjenneberg, S. (1977) Nutritional effect on weight
gain and winter survival of reindeer calves (Rangifer tarandus tarandus). Scientific
Reports of the Agricultural University of Norway 56:1-12.
Jacobsen, E., Hove, K., Bjarghov, R. S., Skjenneberg, S. (1981) Supplementary
feeding of female reindeer on a lichen diet during the last part of pregancy. Effects
on plasma composition, milk production and calf growth. Acta agriculturae
Scandinavica 31:81-86.
Kojola, I., Helle, T., Niskanen, M., Aikio, P. (1995) Effects of lichen biomass on
winter diet, body mass and reproduction of semi-domesticated reindeer Rangifer t.
tarandus in Finland. Wildlife Biology 1:33-38.
Nilsson, A., Danell, ร., Murphy, M., Olsson, K., ร
hman, B. (2000) Health, body
condition and blood metabolites in reindeer after sub-maintenance feed intake and
subsequent feeding. Rangifer 20:187-200.
Rรถnnegรฅrd, L., Forslund, P., Danell, ร. (2002) Lifetime patterns in adult female
mass, reproduction and offspring mass in semidomestic reindeer (Rangifer
tarandus tarandus). Canadian Journal of Zoology 80:2047-2055.
Wiklund, E., Johansson, L., Malmfors, G. (2003) Sensory meat quality, ultimate
pH values, blood parameters and carcass characteristics in reindeer (Rangifer
tarandus tarandus L.) grazed on natural pasturs or fed a commercial feed mixture.
Food Quality and Preference 14:573-581.
Wiklund, E., Pickova, J., Samples, S., Lundstrรถm, K. (2001) Fatty acid
32 R
composition of M. longissimus lumborum, ultimate muscle pH values and carcass
parameters in reindeer (Rangifer tarandus tarandus L) grazed on natural pasture or
fed a commercial feed mixture. Meat Science 58:293-298.
(Oral presentation.)
17
Twenty years of impact of the Chernobyl accident on reindeer
management and meat production in Sweden and Norway.
B. ร
hman
1
and L. Skuterud
2
1
Swedish University of Agricultural Sciences, Reindeer Husbandry Unit, Uppsala,
Sweden
2
Norwegian Radiation Protection Authority and Norwegian Reindeer Husbandry
Administration, Norway
The accident in 1986 at the Chernobyl nuclear power plant caused serious impact
not only locally but also far away from the reactor. Large parts of the reindeer herding
area in Sweden and Norway were severely affected by radioactive fallout. Radioactive
caesium (
134
Cs and
137
Cs) proved to cause a main problem. As an effect of the reindeer
diet, normally containing a large proportion of ground lichen during wintertime, levels
of radiocaesium in reindeer meat from many areas raised to far above what was
accepted for human consumption. This initiated extensive monitoring and control, as
well as a number of different measures to reduce the radiocaesium levels in reindeer
before slaughter.
The levels of radiocaesium in reindeer vary typically with season as an effect of
the seasonal changes in the reindeer diet (Fig. 1). Lichens absorb radionuclides
directly from air and precipitation and get more contaminated than e.g. grass, which
absorbs radiocaesium from the ground. Lichens are eaten predominantly in winter,
whilst the summer diet of reindeer consists mainly of grass, leaves and annual herbs.
The first year after the Chernobyl accident, 78 per cent of the reindeer meat
produced in Sweden was destroyed because of too high levels of radiocaesium. The
highest allowed activity concentration of
137
Cs in most food products at that time was
300 Bq per kg fresh weight. This was later changed to 1500 Bq per kg for reindeer,
game and some other โwildโ products. Consequently, considerably less reindeer meat
(29 per cent) had to be destroyed the following year. In Norway, the intervention level
was first set to 600 Bq per kg (the combined activity of
134
Cs and
137
Cs), but in late
November 1986, this was changed for reindeer meat and game to 6000 Bq per kg. If
the lower level had been kept, 85 per cent of the reindeer meat in Norway would have
been destroyed, while increasing the level reduced condemnation to 27 per cent of the
total production. Since 1994, the intervention level in Norway has been 3000 Bq per
kg.
R 33
0
5000
10000
15000
20000
25000
30000
0 365 731 1096 1461 1826 2192 2557 2922 3287 3653 4018 4383 4748 5114 5479 5844 6209 6575 6940 7305
Bq/kg in muscle
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
Figure 1.
137
Cs in freely grazing reindeer from one herding district, Vilhelmina norra,
in northern Sweden. Each point is the average of data from at least 10 reindeer
measured at the same sampling occasion (meat sampling at slaughter, or live reindeer
monitoring). The reindeer received no feed or caesium binder.
The need for counter measures was obvious in both Norway and Sweden at an
early stage. Already during the first winter after the accident, some reindeer herders
fed their animals uncontaminated feed prior to slaughter. Caesium is relatively mobile
in the body of animals and it is excreted via urine and faeces. If the reindeer has no
intake of radiocaesium, the levels in their bodies is reduced to half the original level
in about two to three weeks (ร
hman, 1996; Skuterud et al., 2004). Feeding has been
a common counter measure to reduce radiocaesium in reindeer before slaughter in
many areas. Compounds that bind caesium in the gut and prevent its absorption into
the body were commonly used during the first years, to enhance the effect of feeding
or to reduce the contamination of freely grazing animals (e.g. Hove & Hansen, 1993;
ร
hman, 1996), and salt licks containing ammonium-iron-hexacyanoferrate (AFCF)
are still used in some reindeer grazing areas in Norway.
Another common counter measure has been the adaptation of slaughter time to the
seasonal variation of radiocaesium. The large part of the reindeer slaughter usually
takes place from November until January, with some additional slaughter later in
winter. By slaughtering earlier in the autumn, contamination of reindeer from the
lichen pasture can be avoided. It was not possible to use this measure in areas with the
highest ground depositions, where reindeer were too contaminated already in early
autumn. As the levels in reindeer declined, early slaughter however gradually replaced
feeding as the most effective and convenient counter measure. However, during the
last years, seasonal variation in radiocaesium in reindeer have been less significant,
and in a number of Norwegian reindeer herding districts there is no difference in
34 R
contamination levels in early autumn compared to winter (Skuterud et al., 2005).
Furthermore, in years when fungal fruit bodies are abundant, radiocaesium levels in
reindeer may be higher in autumn than winter.
There was a clear decline the levels of radiocaesium in reindeer from year to year,
independently of counter measures, shown already a few years after the accident
(Hove et al., 1992; ร
hman & ร
hman, 1994). In Sweden, the rate of decline from 1986
until 2006 corresponds to an effective half-life of 6.2 years at average (recent
estimations using data from 20 out of 52 Swedish reindeer herding districts). The
decline was considerably faster during the first years after the accident than during
later years (corresponding to and effective half-life of 3.6 years for the first ten years,
and 9.6 years for the following ten-year period). In some reindeer herding districts in
Norway (Skuterud et al. 2005), as well as in some of the Swedish herding districts, no
observable decline can be observed during the later ten years.
When the Chernobyl accident happened, there was some radiocaesium
contamination left originating from nuclear bomb tests 20-40 years earlier. It seems
that this โoldโ radiocaesium decline at a considerably slower rate in reindeer than does
caesium originating from the Chernobyl accident (ร
hman et al. 2001). An explanation
to a slower decline with time could be that radiocaesium is gradually removed from
the lichen carpet, but that it still remains in the ecosystem and is available for root
uptake by other parts of the vegetation.
In 2006, 20 years after the accident, the activity concentrations of
137
Cs in reindeer
are below 10 per cent of the initial values. Nevertheless in some regions they are still
above the national intervention levels. Very little reindeer meat is however destroyed
because of too much radiocaesium (less than one per cent during the last ten-year
period). Instead, counter measures like altering the time of slaughter and clean feeding
are used. In Sweden 10โ15 per cent of all slaughtered reindeer are fed to reduce the
contamination. Due to the higher national intervention limit, early slaughter is now the
most common counter measure in Norwegian reindeer herding, affecting about 3โ8
per cent of the reindeer that are slaughtered. Herders are compensated for costs related
to counter measures aimed at reducing radiocaesium levels of reindeer. Presently, the
yearly cost for control and counter measures reaches about one million euro in Sweden
and 300,000 to 400,000 euro in Norway. Although the reindeer herders are
economically compensated, management practises have had to be changed, which
affects reindeer management and the traditional life of the Saami reindeer herders at
large. With the current national intervention levels, we predict that it will take at least
another 10 o 20 years before the impact of the Chernobyl accident on reindeer
husbandry is no longer a problem.
References
ร
hman, B. (1996) Effect of bentonite and ammonium-ferric(III)-
hexacyanoferrate(II) on uptake and elimination of radiocaesium in reindeer.
Journal of Environmental Radioactivity 31:29-50.
ร
hman, B., ร
hman, G. (1994) Radiocesium in Swedish reindeer after the
Chernobyl fallout: seasonal variations and long-term decline. Health Physics
66:503-512.
R 35
ร
hman, B., Wright, S. M., Howard, B. J. (2001) Effect of origin of radiocaesium
on the transfer from fallout to reindeer meat. The Science of the Total Environment
278:171-181.
Hove, K., Garmo, T. H., Hansen, H. S., Pedersen, ร., Staaland, H., Strand, P.
(1992) Duration and variation in radiocesium content of products from domestic
animals grazing natural pasture. Report from the Norwegian Agricultural Advisory
Service 13:256-268 (in Norwegian).
Hove, K., Hansen, H. S. (1993) Reduction of radiocesium transfer to animal
products using sustained release boli with ammoniumiron(III)-
hexacyanoferrate(II). Acta veterinaria Scandinavica 34:287-297.
Skuterud, L., Pedersen, ร., Staaland, H., Rรธed, K. H., Salbu, B., Liken, A., Hove,
K. (2004) Absorption, retention and tissue distribution of radiocaesium in reindeer:
effects of diet and radiocaesium source. Radiation and Environmental Biophysics
43:293-301 (erratum p. 313).
Skuterud, L., Gaare, E., Eikelmann, I. M., Hove, K., Steinnes, E. (2005) Chernobyl
radioactivity persists in reindeer. Journal of Environmental Radioactivity 83:231-
252.
(Oral presentation.)
18
Economic sustainability of farmed venison production in the UK.
M. H. Davies and D. G. Chapple
ADAS UK Ltd, Rosemaund, Preston Wynne, Hereford, HR1 3PG, UK
Deer farming in the UK has maintained a steady state during the last five years.
Recent changes in agricultural support following the Mid-Term Review (MTR) could
have significant implications for deer farming. A desk study was undertaken in 2005
to better understand the economics of farmed venison production compared with other
red meats. Three systems of production were considered; (i) lowland deer units
producing finished animals; (ii) upland deer units producing weaned progeny for
others to finish, and (iii) specialist deer finishing units.
Gross margins achievable from lowland deer farming (ยฃ368 per hectare) now look
more attractive, particularly compared to suckler beef enterprises where subsidy has
historically made up 70-75% of gross margins. However sheep farming, which has
received only 30% of its gross margins from subsidies, still fares better. Gross margins
from upland deer farming are unattractive because of the weak market demand for this
class of stock. In comparison, upland suckler beef systems are facing a difficult future,
but sheep still remain the most attractive option for the uplands. Returns from
36 R
finishing deer units look particularly attractive at ยฃ740 per hectare.
Farmed deer, show a slight negative net margin (after deducting fixed costs) of ยฃ6 per
hind on lowland units, similar to average sheep flocks, but significantly better than
suckler beef. By boosting output through more efficient feeding and pasture
management, net margins approaching those being achieved by the top sheep flocks
should be achievable. However, upland deer units with net margins of-ยฃ42 per hind
are clearly unsustainable. For finishing deer calf enterprises, a positive net margin of
ยฃ30 per stag is achievable and presents a potential opportunity for those lowland
livestock producers considering diversification.
Net margin per kilogram of product is the most important measure of economic
sustainability. Receipts per kg of live weight sold from the three livestock enterprises
show contrasting figures, with finished deer at ยฃ1.75 per kg about 50% higher than
beef at ยฃ1.00-1.15 per kg and lamb at ยฃ1.15 per kg. The high variable costs incurred
in deer farming, together with the very high fixed costs per kg of live weight, lead to
negative margins in both lowland and upland herds (-8p and -113p per kg live weight
respectively). Upland deer production seems completely unsustainable unless calf
selling prices rise substantially, but the significantly higher net margins being
achieved by specialist deer finishing units (30p per kg live weight) are attractive.
(Oral presentation.)
19
Public perception of deer management and control strategies.
M. I. Malins
University of Bath, International Centre for the Environment, Bath, UK
Countryside management policy in the UK now favours wider public access to the
countryside, supported by forestry policy, woodland grant schemes and legislation
promoting access to open country which may bring about potential conflicts with deer
management practice.
UK silvicultural policy has successfully brought about an expansion of forest and
woodland cover, rising from 5.8 % in 1947 to 11.6 % in 2005. This land use change
has favoured the expansion of the wild deer population (Fuller & Gill, 2001) with
apparent impacts on farm crops and trees (Mayle, 1999) wildlife habitats (Cooke,
1995) and disruption to food webs (Kirby, 2001).
In addition to environmental impacts of deer, there are additional public health
concerns regarding bovine tuberculosis, the spread of Lyme disease and an increase
in deer-related road traffic accidents; which has prompted the calls for increased
culling of deer, including the wider use of amateur hunters.
A view of the public perception of deer management was obtained through a UK
wide survey undertaken in 2004/5 targeted at recreational users of the countryside.
Responses were received from a cross-section of socio-economic groups who
provided information on the type, frequency and location of their activities undertaken
R 37
in the countryside.
Public agreement with the principle of deer management, preferred strategies for
reducing road accidents; favoured deer control methods, the sale of venison and the
role of state agencies are reported. Additional evaluation of the respondents' awareness
of deer management, perceived compatibility of public access and shooting, risk
reduction strategies and preference for UK or EU training standards are also provided.
References
Cooke, A. (1995). Muntjac damage in woodland. Pages 1214 in Enact managing land
for wildlife. 3 (3). English Nature. Peterborough.
Fuller, R.J. and Gill, R. (2001) Ecological impacts of increasing numbers of deer in
British woodland. Forestry 74: 193-199.
Kirby, K. (2003) Deer and biodiversity action plan targets. Pages 1823 in Goldberg,
E., editor. Proceedings of the Future for Deer Conference. English Nature,
Peterborough.
Mayle, B. (1999). Managing Deer in the Countryside. Forestry Commission. Practice
Note FCPN6.
(Oral presentation.)
20
Preferences of red deer for subtropical pasture species.
G. M. Dryden and K. J. Whelan
School of Animal Studies, The University of Queensland, Gatton, QLD 4343, Australia
The subtropical Australian deer industry is pasture-based but there is very little
information on the preferences of deer for the pasture species present in this region
and which form the industrys main food resource. An investigation into the diet
selected by red deer (Cervus elaphus) grazing tropical pastures was conducted at the
University of Queenslands Gatton Deer Research Unit in the summer of 2003. Twenty
mature hinds (5 and 6 years old, 102.9 ยฑ 10.10 kg) were allocated to one of two
paddocks containing 22 plant species. Predominant species were the forb Cyperus
rotundus, the native grasses Eleusine indica and Sporobolus elongatus, and the
naturalised grasses Chloris guyana, Cynodon dactylon, Cynodon nlemfuensis and
Pennisetum clandestinum. Faecal alkanes were used to investigate the diet
composition in five consecutive weeks. The main species in the selected diets were
C. dactylon (overall mean ยฑ se, 38.5 ยฑ 9.22%), C. nlemfuensis (6.5 ยฑ 0.81%) and C.
rotundus (53.9 ยฑ 1.15%). The other species were largely avoided. There were small
differences in the diets selected by the deer in the two paddocks: for C. nlemfuensis
those in Paddock 1 ate 5.3% v. 7.7% for Paddock 2 (P=0.0368), and for C. rotundus
the Paddock 1 diet contained 57.4% v. 50.2% in Paddock 2 (P=0.0074). The
38 R
differences for C. nlemfuensis and C. rotundus reflected the abundances of those
species in the paddocks. Diet preferences evolved throughout the experiment:
selection for C. dactylon and C. rotundus increased after Week 1, while that of C.
nlemfuensis was reduced. This behaviour suggests that the deer avoided C.
nlemfuensis as it matured. C. dactylon may have been consumed because it was the
predominant species in both paddocks. The results show that C. dactylon, which is a
common component of southeast Queensland pastures, is a suitable species for deer
farms, and that C. rotundus, which is often considered a weed, may be more valuable
than generally thought.
(Oral presentation.)
21
Evaluation of forage herbs for farmed red deer: feeding value and
trace elements.
S. O. Hoskin, P. R. Wilson, M. Ondris, and A.-H. Bunod
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
This paper summarises three studies aiming to: compare the copper, vitamin B12 and
selenium status of young farmed red deer grazing perennial ryegrass-based pasture
(Lolium perenne), chicory (Cichorium intybus) or plantain (Plantago lanceolata) in
spring; contrast the feeding value of perennial ryegrass-based pasture with chicory and
plantain in spring and plantain in autumn; and determine the effect of grazing
perennial ryegrass-based pasture alone, or with intermittent grazing of plantain from
one month pre-partum to one month post-partum on copper and vitamin B12 status of
calves. Feeding value, determined by live weight gain, of deer grazing chicory was
greater than for deer grazing plantain and pasture in spring and feeding value of
plantain was greater than pasture in autumn. Grazing chicory significantly enhanced
the copper status of weaner deer in spring compared with both pasture and plantain.
Grazing plantain significantly enhanced the copper status of weaners in autumn but
not weaners or younger calves in spring and early summer, enhanced the vitamin B12
status of weaners in autumn and spring and calves in early summer, and boosted the
selenium status of weaners during both spring and autumn. This study has highlighted
the complementary role of forage herbs in achieving higher growth rates and
maintaining trace element status of young farmed deer.
(Oral presentation.)
R 39
22
Habitat Utilization by Himalayan Musk Deer (Moschus
chrysogaster), Sambar ( Cervus unicolor) and Barking Deer
(Munitacus muntjac) at Kedarnath Wildlife Sanctuary, Western
Himalaya.
S. Sathyakumar, S. N. Prasad, G. S. Rawat, and A. J. T. Johnsingh
Wildlife Institute of India, P.O. Box 18, Chandrabani, Dehradun 248 001, India
We investigated the habitat utilization by Himalayan Musk Deer (Moschus
chrysogaster), Sambar (Cervus unicolor) and Barking Deer (Munitacus muntjac) from
1989 to 1991 in a 20 Km2 intensive study area that had 10 habitats and altitude
ranging from 1700m to 3680m, with diverse aspect and slope categories at the
Kedarnath Wildlife Sanctuary, Uttaranchal, India. For Musk deer and Sambar in the
subalpine zone, we estimated density using the silent drive count method and collected
habitat use data by systematic searches. We estimated encounter rates, density
estimates and habitat use by Barking deer and Sambar in the Temperate zone using
line transects (N=5; L=1 to 4.8 km). In the subalpine forests, Musk deer and Sambar
density estimates were 3.7/km
2
and 0.9/km
2
respectively. The encounter rates and
density estimates for Barking deer and Sambar in the temperate forests were 0.37 and
0.54/km walk; and 2.9 groups/km
2
(mgs=1.6) and 2.5 groups/km
2
(mgs = 1.9),
respectively. Musk deer (n=52) mostly used the 3,000-3,300m altitude range, southern
and south western aspects and 41o- 60o slope categories. It mostly occurred in sites
that had 26-50% tree cover; 1-50% shrub cover, and 1-25% ground cover. Sambar
(n=75) mostly used the 2,100-2,400m altitude range, north eastern aspect and 31o-
40o slope categories. It mostly occurred in sites that had 1-50% tree, shrub, and
ground cover categories. Barking deer (n=117) mostly used the 1,700-2,000m altitude
range, north eastern aspect and 31o- 40o slope categories. It mostly occurred in sites
that had 26-75% tree cover; 1-50% shrub cover, and 1-50% ground cover. Musk deer
used the alpine scrub habitat more than its availability, Sambar used the Temperate
oak-fir-maple habitat more than its availability, and Barking deer used the Temperate
oak-alder habitat more than its availability and all other habitats were used by these
three species in proportion to their availability. The differential use of altitude by these
three species was the major reason for their ecological separation. Although the musk
deer and barking deer are similar in body size and being small solitary forest
ruminants, they are considered allopatric whereas Sambar with its large body size was
more sympatric to barking deer.
(Oral presentation.)
40 R
23
Feeding habits of red deer in Hungarian forested and agricultural
areas.
K. Katona, L. Szemethy, K. Mรกtrai, N. Bleier, and Sz. Orosz
St. Istvรกn University, Department of Wildlife Biology and Management, Pรกter Kรกroly
u. 1., Gรถdรถllล 2103, Hungary
The red deer (Cervus elaphus L. 1758) is a very important large herbivore in the
Hungarian fauna. This species has high ecological and economical impact on its
environment. Department of Wildlife Biology and Management of St. Istvรกn
University has carried out long-term studies on the diet composition and feeding
characteristics of red deer in Hungary. The results, however, are often contradictory
with earlier knowledge and beliefs about feeding ecology of red deer. Our
investigations conducted in several forested and agricultural habitats of Hungary
revealed the following important facts: 1.In forested habitats red deer dominantly
consumed the different browse species of the understory vegetation.
2.In agricultural habitats red deer preferred available browse species (along canals, or
in forested patches). Cultivated plant species (mainly maize) were not preferred, their
proportion in the diet was very low. 3.The proportion of grasses in the forest diet was
also very low, its consumption was much higher in agricultural areas, but red deer did
not show preference to grasses. 4.Red deer diet was highly variable and well-adapted
to the actual food supply of different habitats. Some species were highly preferred in
all areas (Robinia pseudoacacia, Rubus spp., Sambucus spp., Cornus sanguinea, Salix
spp.), but many other browse species also appeared in the diet. 5.Quality (crude
protein/crude fiber ratio) of the preferred diet components was higher, than the other
species. 6.Plant species available in hunting grounds or provided as winter
supplemental food were found in very low proportion in the diet of red deer. One
exception was Medicago sativa - with high protein content - , which was regularly
consumed. Our results can be supported with ecophysiological constraints of red deer.
The species is an intermediate feeder, which is able to consume a higher proportion
of grasses, but it has to eat a high proportion of good quality food. Problems of game
damages in forested and agricultural areas can be decreased by considering our results
on the feeding biology and behaviour of red deer.
(Oral presentation.)
R 41
24
Detection of needles: tool for evaluation of diet quality in wild
ruminants.
J. Kamler, M. Homolka, M. Heroldovรก, M. Baranฤekovรก, and J. Prokeลกovรก
Institute of Vertebrate Biology, Academy of Sciences of the Czech Republic, 603 65
Brno, the Czech Republic
Norway spruce needles are a wide-spread food resource and their utilisable biomass
exceeds the needs of herbivores. Needles seem to be a generally ignored food source
that is consumed at the last place especially during severe winters with deep snow
cover. Aim of our study was to examine the possibility of evaluation of needles in the
diet as an indicator of quality of diet of free living large herbivores. We verified low
quality and dietary preference of spruce needles in comparison to other ungulate diet
resources in winter. Consummation of spruce needles can be used as a good indicator
of quality of the environment and suitability of herbivore density in most localities in
the central Europe. We developed easy applied technique using near infrared
spectrophotometer to determine the content of spruce needles in the faeces.
(Oral presentation.)
25
Environmental factors affecting Scots pine debarking by red deer
in south-western Poland.
J. Borkowski and P. Nasiadka
Forest Research Institute, Section of Forest Ecology and Wildlife Management,
Sekocin-Las, Poland
Scots pine debarking by red deer was studied in sixth (1999) and seventh (2000) years
after the large (10,000 ha) forest fire in south western Poland. The main aims of the
study were to: i) compare debarking in artificially regenerated prethickets with that in
stands regenerated naturally; ii) check if security cover conditions (dependent on the
height of prethickets) influence debarking intensity and iii) check if debarking of the
prethickets depends on the tree density.
There was no significant difference in the debarking intensity (% of debarked trees)
between artificially and naturally regenerated stands neither in 1999 nor in 2000.
When similar comparisons were made controlling the age (and at the same time the
height) of artificially regenerated stands (majority of naturally regenerated thickets
were of the same age), in 1999 naturally regenerated thickets were debarked with
similar intensity as the plantations of the same age. However, younger pine plantations
were debarked less and older plantations more than the stands coming from natural
42 R
regeneration. In 2000 (when all the thickets got high enough to provide deer with
favorable cover) all the plantations were damaged with the same intensity as naturally
regenerated stands. In general, debarking in pine plantations was positively correlated
with their age but only up to eight years old and further increase in age did not
influence debarking intensity. The relation between percent of debarked trees and the
height of artificially regenerated pine stands depended on their age. In younger
prethickets debarking intensity was correlated with their height, while in older ones
the relation disappeared. It can be concluded that security cover is important
determinant of debarking intensity of Scots pine prethickets. Tree density had no
significant influence on debarking intensity in the plantations nor in the naturally
regenerated stands.
(Oral presentation.)
26
New technique for estimation Cervidea hiding cover.
A. J. Ferreira and A. M. Oliveira
Instituto Ambiente e Vida - Universidade de Coimbra. Largo Marques do Pombal.
3004 -517 Coimbra. Portugal.
Hiding cover is a key factor for habitat selection by Cervidae, and its knowledge is a
matter of great concern to wildlife management. Usually evaluating of the animal
hiding cover is performed by measure the vegetation horizontal capacity of cover one
target. These techniques present some difficulties due to observation bias and field
resources. A new horizontal cover measurement method was tested and compared to
most commonly used techniques for cover valuation. It is based on digital analysis and
edition of images of wide sampling areas. It proved to be an effective technique with
no different results from other methods, as well as easier and faster to manage in the
field, allowing higher number of samples with similar effort. Besides, it proved to be
more sensitive to extreme cover situations, resulting into more real estimations.
Keeping a digital record of the samples is another advantage of our method, because
posterior analysis becomes independent of field sampling.
(Oral presentation.)
R 43
27
Calving sites fidelity in free-ranging moose.
J.-P. Tremblay
1
, E. Solberg
2
, B.-E. Sรฆther
1
, and M. Heim
2
1
Department of Biology, Norwegian University of Science and Technology (NTNU)
2
Norwegian institute for nature research (NINA)
Northern ungulates are generally considered to exhibit fidelity to their previous
calving sites conditionally to the spatial scale considered. Much of our knowledge
about the tendency of these animals to return to the same area to give birth is drawn
from open-dwelling species as caribou (Rangifer tarandus). Estimation of calving sites
fidelity is however notoriously difficult to achieve for inconspicuous forest ungulates
as moose (Alces alces) and deer. Our objective was to quantify fidelity of moose to
their calving sites and if so to determine how individual life history influence patterns
of site fidelity. We used long-term monitoring (1992-2005) from an isolated island
population of moose in northern Norway (Vega, Nordland county) consisting of a
dataset of 31 moose cows with at least two, and up to nine known calving site
locations. Locations were determined based on intensive radio-locations of marked
females and direct observations of calves in the first few days following birth. We
generated null expectations of fidelity from the distribution of distances between
locations of calving sites from all females in the population comprised within the
home range of each specific female (home range scale) and from all known calving
sites on the 119 km island (landscape scale). We contrasted distances between
consecutive-year locations of calving sites to these expectations using log-linear
regression analysis on the ratio of observed: expected distances. Site fidelity has been
shown to exhibit individual variation based on previous age and previous reproductive
success. Female experience (age, number of offsprings produced and recruited) and
other fitness correlates (body mass) were used to explain the variability in site fidelity
within the population. The effect of the mother fitness on calving site fidelity of her
daughters was also investigate using 26 female-daughter pairs with known calving
locations (from 14 adult females). Results from this study contribute to our
understanding of the effect of this ecological pattern of habitat use on the demographic
stochasticity in moose population.
(Oral presentation.)
44 R
28
Spatio-temporal distribution of white-tailed deer relative to
prescribed burns on rangeland in south Texas, USA.
M. G. Meek
1
, S. M. Cooper
2
, M. K. Owens
2
, and A. L. Wappel
2
1
Department of Wildlife and Fisheries Sciences, Texas A&M University, College
Station, TX, USA
2
Texas A&M University System, Texas Agricultural Experiment Station, Uvalde, TX,
USA
This study is being done to determine the spatial and temporal distribution of white-
tailed deer (Odocoileus virginianus) on prescribed burned areas in a rangeland
restoration project. Overgrazing and fire suppression has left much rangeland in the
United States in poor habitat condition. Prescribed fire is one method used to increase
the range health. Browsing by deer may have a major impact on the regrowth of
vegetation. Three areas of rangeland, each of 40 ha will be burned each year. To
determine habitat use and distribution of deer relative to these burns 3 bucks and 3
does will be netted from a helicopter and fitted with Global Positioning System
(G.P.S.) telemetry collars (Lotek GPS 3300s) for a period of 45 days during each
season. These collars are programmed to take a position fix every hour which limits
problems due to spatial autocorrelation. Six deer were collared in August 2005, prior
to implementing the prescribed burns. This preliminary data showed that the future
treatment areas were under-utilized by deer. The next three collaring seasons will
follow response of deer to the burns as the vegetation matures. At this point, the first,
second, and third trials have been completed. The second trial was approximately one
month after burning and the third was at spring green up. Thus far, the data indicates
that bucks use both burned and untreated areas but does tend to avoid the newly
burned areas. Whether this is due to reduced browse, loss of escape cover, or natural
weariness are unknown at this point. The second trial was done during the breeding
season which could affect movement patterns, especially for bucks. Analysis of the
spring trial is ongoing. Geostatistical analysis will be applied to spatial and temporal
data collected by the G.P.S. collars to determine spatial dependence of deer
distributions, on vegetation and burned areas. The Animal Movement extension in
ArcView 3.2 and Hawths Tools extension in ArcGIS 9.1 will be used with ANOVA
to relate habitat use with vegetative characteristics.
(Poster presentation.)
R 45
29
Sexual segregation and differences in quality of diet in white-tailed
deer (Odocoileus viginianus mexicanus) in a tropical dry forest in
Mexico.
A. Buenrostro
1
, S. Gallina
2
, and G. Sรกnchez-Rojas
3
1
Universidad del Mar,Carrera de Zootecnia,Puerto Escondido, Oaxaca, Mexico;
2
Instituto de Ecologรญa, A.C. km 2.5 Carretera Antigua a Coatepec No. 351,
Congregaciรณn el Haya, Xalapa, Veracruz, CP 91070, Mรฉxico
3
Laboratorio de Conservacion Biologica,Centro de Investigaciones Biologicas,Area
Academica de Biologรญa,Instituto de Ciencias Basicas e Ingenieria Universidad
Autonoma del Estado de Hidalgo,Apostado Postal 69 Plaza Juarez, Pachuca, Hidalgo
42000 Mexico
Sexual segregation in deer can be seen at food level, as dietary composition reflects
diverse biological aspects, demonstrating intersexual differences. Some authors
suggest that females select higher quality diets, which is attributed to differences in
body size and reproductive strategies. The present study evaluated the diet quality, as
well as temporary and intersexual variations in the white-tailed deer through non-
invasive fecal indexes such as the nitrogen fecal content (NF). The objective was to
test the following predictions: a) NF concentrations will be higher in females, and b)
differences will exist between reproductive and non-reproductive periods. The study
was conducted in a tropical dry forest in the state of Morelos, Mexico. Five sample
collections were obtained from January through May, 2004. A total of 113 groups of
fecal samples were analyzed individually and in duplicate, of which 63 belonged to
females and 50 belonged to males (the pellet morphometry was used to distinguish sex
by the fuzzy set method) at two reproductive periods: one corresponding to mating
season, (when both sexes are together) and the second, the non-reproductive period
(when the sexes are separated). The period between sampling did not exceed 30 days,
to avoid loss of fecal nitrogen. The percentages of NF between both sexes showed
significant differences (F = 5.88; g.l. = 1; P = 0.017). NF concentrations in fecal
groups belonging to females was evidently greater; however, no significant difference
was found between reproductive and non-reproductive periods (F = 2.48, g.l. = 1,111,
P = 0.1181). These results show that both sexes during mating season are occupying
better habitats with high quality food, while in dry season females increase their
quality of diet despite adverse conditions. This confirms that sexual segregation can
be reflected at the level of diet.
(Poster presentation.)
46 R
30
Sex comparison of linear body measures of growing red deer calves
(Cervus elaphus hippelaphus).
B. Dmuchowski
1
, M. Snochowski
2
, and A. Krzywiลski
3
1
Research Station, The Witold Stefaลski Institute of Parasitology, PAS, Kosewo Gรณrne,
11-700 Mrgowo, Poland;
2
The Kielanowski Institute of Animal Physiology and Nutrition, PAS, 05-110
Jablonna, Poland;
3
Wild Life Park, Kadzidลowo, 12-220 Ruciane-Nida, Poland.
The aim of the present study was to investigate whether the sex differences affects the
body shape of red deer calves below the age of 16 months. The study was performed
on 186 female and 166 male calves born between 1998 and 2005 in Kosewo Research
Station (53ยฐ42โ N, 21ยฐ38โ E). About 90 % of animals originated from natural
population in Poland and less then 10% were 12.5 % Wapiti or Maral hybrids. All
calves were kept from birth until late autumn together with hinds on natural pasture.
Prior the winter they were separated from hinds and fed the diet consisted from
mixture of oats or barley and ground soy bean with free access to hay or haylage in
amounts calculated for daily intake of 1.5 g total protein and 0.16 MJ metabolic
energy per kilogram of body weight. After wintering all calves were put to pasture
again. Since there was no significant effect of the year on analysed parameters the data
were calculated together with respect to sex and age. The life weight (LW; 1372
records) increased with age in the sex specific manner, and were fitted to quadric
equations (p<0.001) Y= -0.0343x
4
+1.183x
3
-13.73x
2
+65.77x-38.64 for males and Y=
0.0252x
4
+0.904x
3
-11.00x
2
+55.14x-30.67 for females, where Y is LW (kg) and X is
month of life (3 to 15). The linear body measures analysed (496 records/parameter)
included body length (BL), head length (HL), lower jaw length (LJ), length of front
pastern (FP) and back pastern (BP), and chest depth (CD). The changes of all
measures were found to be strictly related to LW and were not dependent on sex. They
were fitted to following quadratic equations (p<0.001) Y=aX+b, were Y is respective
parameter (cm) and X is LW (50-150 kg): BL = 0.461X+123.8; HL = 0.140X+25.2;
CD = 0.131X+26.6; BP = 0.106X+31.2; LJ = 0.097X+18.4; FP = 0.065X+21.5. Based
on linear body measures analyzed we conclude that no sexual differentiation of body
shape occur in red deer calves during the first 16 months of life and parametric
description of accumulated data may serve as reference for comparative studies.
(Poster presentation.)
R 47
31
The influence of management system of farmed fallow Deer (Dama
dama) on selected production traits during winter season.
B. Dmuchowski
1
, J. Starz
2
, A. Demiaszkiewicz
3
, and R. Niลผnikowski
2
1
Research Station, The Witold Stefaลski Institute of Parasitology, PAS, Kosewo Gorne,
11-700 Mrgowo, Poland;
2
Warsaw Agricultural University (SGGW), Faculty of Animal Sciences,
Nowoursynowska 166,02-787 Warsaw, Poland;
3
The Witold Stefaลski Institute of Parasitology, PAS,Twarda 51/55, 00-818 Warsaw,
Poland.
The aim of this study was to determine optimum method of winter management
system of farmed fallow deer (Dama dama). The investigation was conducted on
group of 73 calves both sexes. In December 2004 animals were drenched (injection
of 1% ivermectin), weighted and divided into two groups. The first group was held in
half-open shed during winter season. The second group of calves was wintering along
with hinds on winter pen. In May 2005 calves became weighted again. From animals
(before and after wintering), chosen at random, was taken samples of faeces, to make
parasitological explorations. Analysis of weight exhibited, that calves, which were
hold in the shed, had bigger increase of body weight. The parasitological explorations
proved that drenching was more effective in group of calves held in the shed, without
contact with hinds.
(Poster presentation.)
32
Deer home range overlap and habitat heterogeneity in Northeastern
Mexico.
J. Bello, S. Gallina, M. Equihua, and N. Corona
1
Divisiรณn Acadรฉmica de Ciencias Biolรณgicas, Universidad Juรกrez Autรณnoma de
Tabasco, km 0.5 Carretera Villahermosa-Cรกrdenas, Villahermosa,Tabasco,
CP 86039, Mรฉxico
2
Instituto de Ecologรญa, A.C. km 2.5 Carretera Antigua a Coatepec No. 351,
Congregaciรณn el Haya, Xalapa, Veracruz, CP 91070, Mรฉxico
The knowledge about the use of habitat by animals and how it change with
environmental conditions is important for management programs. We analyzed deer
home range overlap between sexes (13 females and 10 males), seasons and years, and
the relationship with habitat heterogeneity using radiotracking during 1994 to 1998.
The data were grouped in three physiological seasons: 1) Reproductive or breeding
48 R
(November to February), 2) Posreproductive or gestation (March to June) and 3)
fawning (July to October). This research was carried out at the San Francisco Ranch
located in Northeastern Mexico. The area is an 1000 ha enclosure for deer . An
intensive water management program operate in the area (3 dams and 32 water
troughs). The climate is semiarid, with a mean annual temperature of 21C. Annual
rainfall averages less than 400 mm, with notable variation. The vegetation is a
xerophyllous brushland. The habitat heterogeneity was obtained within cells (1000
m
2
), taking into account the total borders and number of habitat types. Home range
was estimated with the Minimum Polygon Convex model. Overlap was obtained
between females, males and males-females, for each season and year. ANCOVA was
used to know significant differences between sexes, seasons and years. More than
70% of overlap was obtained during post-reproductive season (dry season) for the
three sex interactions. The less overlap occurred between females during fawning
season, mainly in dry years, so this demonstrate that females prefer isolation during
fawning, maybe to secure resources for their fawns. There was a significant positive
correlation between habitat heterogeneity considering borders and number of deer
locations. The number of habitat types by cell varied from one to five, with the highest
number of deer locations in cells with three and four habitat types and the least with
one habitat type. Deer prefer habitat heterogeneity because these areas had higher
resources diversity for cover their requirements.
(Poster presentation.)
33
Influence of ranging strategy on home range size: red deer hinds in
a forest-agriculture habitat.
Zs. Birรณ, L. Szemethy, and K. Katona
St. Istvรกn University, Department of Wildlife Biology and Management, Gรถdรถllล 2103,
Hungary
Radiotelemetry studies of 27 red deer (Cervus elaphus Linnรฉ, 1758) hinds in a forest-
agriculture habitat between 1993 and 2000 revealed two stable alternative ranging
strategies. Shifters relocated their home range from the forest to the agricultural area
during summer. Residents stayed in the forest throughout the entire year. Because of
these clear differences between ranging behaviours and the two habitats substantial
differences were expected in the home range size between the two groups. Three
theories were taken into consideration: body-size, habitat-productivity and habitat-
heterogeneity hypotheses, all of which predict larger seasonal home ranges in the case
of a suboptimal ranging strategy. Consequently, summer and winter home range sizes
were compared between strategies and seasons. Both entire home ranges and core
areas were significantly larger for both strategies in winter than in summer. There was
no difference between strategies during the summer period. Contrarily, both entire
winter home ranges and core areas of shifters were significantly larger than that of
R 49
residents. Our findings suggest that shifters suffer disadvantages in the forest in
winter. These results could not be clearly explained by any of our hypotheses. We
suggest social factors to be considered for a better understanding.
(Poster presentation.)
34
New project on red deer Cervus elaphus in Sweden.
A. Jarnemo
Grimsรถ Wildlife Research Station, Swedish University of Agricultural Sciences,
Sweden
A new research project on red deer Cervus elaphus was started in Sweden in 2005.
The aim is to produce data that can be used to improve management of red deer in
Sweden. The project will study home range, movement patterns, habitat choice,
reproduction, monitoring methods and management practices. The project uses two
study areas: one in southernmost Sweden (Skรฅne) and one in central Sweden
(Kolmรฅrden).
(Poster presentation.)
35
Mapping of male red deer Cervus elaphus movements in southern
Sweden.
A. Jarnemo
Grimsรถ Wildlife Research Station, Swedish University of Agricultural Sciences,
Sweden
Summer and winter sites used by individually identified male red deer in southernmost
Sweden (Skรฅne) were linked to areas where the stags appeared during the rut.
Individuals were identified on antler characteristics and photo-documented during rut.
Trophies from harvested deer and carcasses were documented as were found cast
antlers. Stags were also photo-documented during summer. From 1970 to 2005 a link
between summer/winter site and rutting ground could be established for 58
individuals. The mean distance between rut documentation and summer/winter
documentation was 15 km, ranging from 2.5 to 47 km. Mapping of male red deer
movements can help to establish the minimum area needed for an effective joint
management and to coordinate harvest and management between rutting grounds and
winter-summer areas of males. The identification of the summer-winter areas should
50 R
also serve as a first step in an attempt to study the causes of sexual segregation of red
deer in the area.
(Poster presentation.)
36
Importance of floodplain forest for deer management.
J. Prokeลกovรก, M. Baranฤekovรก, and M. Homolka
Institute of Vertebrate Biology AS CR, Kvฤtnรก 8, 603 65 Brno, Czech Republic
Floodplain forests represent despite their small size very important forest habitat in the
Czech Republic from point of view of natural conservation. They are characterized by
high biodiversity, which could be decreased by human activity (regulation of rivers,
transformation to agricultural area) as well as by high abundance of big herbivores
(browsing, trampling, bark stripping). Most of the floodplain forests are commercially
managed for timber harvesting so for forestry the presence of deer is because of its
browsing impact on planted seedlings undesirable.
The aim of our study was to estimate the density of red and roe deer and to assess their
impact on biodiversity of floodplain forest ecosystem from point of view of carrying
capacity of this habitat, using mainly the height of browsing impact. Based on these
results we tried to make required conclusions for deer management.
During the research in 2002 โ 2004 we found that the densities of both deer species
are above-averaged (9.6 ind./km
2
and 7 ind./km
2
, red and roe deer respectively) in
comparison with other forests in Czech Republic. In spite of high proportion of
broadleaved trees in their diet (red deer 71%, roe deer 56%), the browsing impact did
not exceed 30% and is relatively low in comparison with other forested areas. The
shrub layer is very well developed and regenerates naturally. On the other hand
browsing is an important threat to artificially planted oak. They have to be fenced.
High carrying-capacity of floodplain forest makes it possible to keep high abundant
populations of red and roe deer. In spite of high density and high proportion of trees
in deer diet, the natural regeneration of the forest and its biodiversity are not
threatened. But high deer density causes problems to forestry and the artificial
plantations of economic tree species have to be protected against browsing. Their
protection by fencing is quite effective and for the present the reduction of deer
abundance is not necessary.
The study was supported by GA AS CR S6093003 and by grant No. LC06073 of the
Czech Ministry of Education.
(Poster presentation.)
R 51
37
Gross composition and protein fractions of milk from fallow deer
(Dama dama).
G. M. Pisani, M. Malacarne, C. S. Soffiantini, P. Franceschi, P. Formaggioni, E.
Piasentier
1
, A. Summer, and P. Mariani
Department of Animal Production, Veterinary Biotechnology, Food Quality and Safety
- University of Parma, Italy
1
Department of Animal Science - University of Udine, Italy
Thirty milk samples were collected from 18 lactating does lambed on June and July.
The lactating herd comprised 12 adult animals older than 3 years (mean age: 42
months) and 6 young animals in their third year of live (mean age: 23 months).
Sampling was carried out in late August 2004 (13 milk samples) and in the same
period of the year 2005 (17 milk samples). Does were reared at the deer research
facility Antonio Servadei (University of Udine), located in Pagnacco, north-east of
Italy. The lactating herd was allowed to graze a pasture dominated by Festuca
arundinacea and daily supplemented on 7 kg of a concentrate feed based on cereal
grain and soybean meal (CP 20.3 % on dry matter basis). During milking, deliveries
were maintained in the does herd. Does were immobilised by means of a crush and 10
I.U. of oxytocin were injected by intramuscular administration to facilitate milk
ejection. After 1-2 minutes, does were handly milked as deep as possible in 10-20
minutes. About 70 mL of milk was collected from each does. On milk samples, the
following parameters were determined: total nitrogen (TN), soluble nitrogen (SN) and
non protein nitrogen (NPN), from which total protein (TN x 6.38), casein ((TN - SN)
x 6.38), true whey protein ((SN-NPN) x 6.38); fat and lactose and ash contents. Dry
matter, total protein, fat, lactose and ash contents (meansd) of fallow deer milk
resulted, respectively, 29.27ยฑ3.13, 8.04ยฑ1.16, 14.92ยฑ2.42, 3.25ยฑ0.27 and 1.08ยฑ0.12
g/100g. Concerning protein fractions, the contents of casein, true whey protein and
NPN x 6.38 resulted, respectively, 6.89ยฑ1.11, 0.79ยฑ0.12 and 0.30ยฑ0.08 g/100g. Csapo
et al. (1987) in a study carried out on milk samples collected in two occasions (15-20
days after dropping and in the second-third month of lactation) from 3 hinds of fallow
deer reared in captivity, reported values clearly lower of dry matter (-33%), total
protein (-14%) and fat (-44%) than those observed here. As far as protein fractions,
compared to our results, the value of casein content resulted lower (-20%), while true
whey protein and NPN x 6.38 contents were similar.
(Poster presentation.)
52 R
38
Estimating in vitro digestibility of wild sika deer (Cervus nippon
yesoensis) in Hokkaido, Japan.
C. Yayota, K. Nishitani, K. Ueda, Y. Yanagawa, Y. Matsuura, M. Suzuki, H. Hata,
and S. Kondo
Hokkaido University, Guraduate School of Agriculture, Nishi 9 Kita 9 Kita-ku
Sapporo-shi, Hokkaido, Japan
In Hokkaido, a northern-most island of Japan, the damage to agriculture and forestry
by sika deer has recently increased. The nutritional information such as intake and
digestibility of wild sika deer is one of the most important information for the
population management and accurate estimation of the carrying capacity, while
determination of digestibility for wild sika deer have been fairy difficult. In this study,
in vitro method was applied, 1) to determine the digestibility of wild sika deer, 2) to
compare them with those of cattle (Bos taurus). In Experimental Farm of Hokkaido
University, four and five wild female sika deer were harvested in November (fall; FA)
and February (winter; WI), respectively. The rumen fluid was collected from these
deer and used to measure in vitro digestibility and gas production immediately.
Additionally, in vitro digestibility using the rumen fluid collected from three cattle
with rumen cannula was measured. To compare in vitro digestibility and gas
production of deer and cattle, dried Lolium perenne (L), leaves of Sasa nipponica (S),
leaves of Carpinus cordata (C) and barks of Acer mono (A), were used. In vitro
digestibility of dry matter (DM), crude protein (CP) and neutral detergent fiber (NDF)
were determined after 48-hour incubation. In vitro gas production was measured at
regular intervals thoughtout 96-hour incubation and calculated the parameters by
curve fitting. Content of CP was 27.3, 16.7, 14.1 and 7.6% of DM, and the NDF
content was 39.5, 65.3, 55.8 and 74.1% of DM in L, S, C and A, respectively. The
digestibility was the highest in L and the lowest in A in both deer and cattle.
Digestibilities of CP and NDF for all plants in deer decreased in WI, especially in C
and A. The asymptotic gas production (ml/gOM) was the highest in L in both deer and
cattle. The fractional degradation rate (/h) in deer was faster in FA than in WI. In vitro
digestibility and gas production parameters were significantly correlated between deer
and cattle. These results suggested that in vitro digestibility and gas production in wild
sika deer could be estimated by those in cattle.
(Poster presentation.)
R 53
39
Comparison of physical condition of two Red deer (Cervus elaphus)
populations.
A. J. Ferreira and R. M. Ramalho
Instituto Ambiente e Vida - Universidade de Coimbra. Largo Marques do Pombal.
3004 -517 Coimbra. Portugal
The body condition of Cervidae is usually an important source of information about
the population performance. In the present study we evaluated the physical condition
of two red deer sub-populations localized in the same climate area (Mediterranean
climate) that are apart of which other by about 30 km. However with the same climate
conditions, the two areas have a distinct management history, manly in the land use.
One population is localized in a characteristic mediterranean habitat, composed mainly
by Quercus spp. and the other in a highly managed conifers forest. Using a
combination of the kidney fat index (KFI) and the bone marrow fat (BMF) we could
compare the performance of both sub-population that have the same genetic origins
and similar management practices. Some considerations about the combined use of
these two indexes and the land management option are address in this work.
(Poster presentation.)
40
Distribution, abundance and management of the two native deer in
Italy.
L. Carnevali, F. Riga, and S. Toso
INFS - National Wildlife Institute, Bologna, Italy
In the last decades the distribution and abundance of roe deer and red deer, as well as
several other Italian ungulates, constantly increased, but this expansion phase occurred
unevenly throughout the peninsula. Aim of this poster is to present the distribution and
abundance of the two Italian native deer and to illustrate the development in their
population dynamics and management strategies starting from 1999-2000 (Pedrotti et
al., 2001). The data were provided by the National Ungulates Database, implemented
by the National Wildlife Institute (INFS), in which all the available information
regarding wildlife censuses carried on by local administrations and protected areas,
as well as harvest plans and annual hunting bags starting from 1996 are recorded. At
present, roe deer (most widely distrib among Italian deer) and red deer occur in the
most of the alpine range, while the distribution in the northern-central Apennines
range is wide for the roe deer and scattered for red deer. In the south, red deer is
present only in two national parks in which re-introduction programs have been
54 R
recently carried out and roe deer in four national parks: in three of them is present the
sub-species Capreolus capreolus italicus, also present in a fenced protected area near
Rome (Castel Porziano) and in a part of central Italy. The Sardinia Island is inhabited
by the red deer sub-species C.e.corsicanus (6,000 animals estimated in 2005), and a
relict population of the ancient Italian species is still present in Ferrara Province with
about 100 animals (Mesola red deer). In 2000, about 336,000 roe deer and 44,000 red
deer were estimated and about 30,000 roe deer and 4,000 red deer have been harvested
in Italy. In a few years, roe deer population have increased of about 25% (420,000
animals estimated in 2005), even if the pattern is not homogeneous in the entire
country, and the red deer population of about 30% (56,000 animals estimated in 2005).
Following the population trends, harvest quotas increased and about 48,000 roe deer
(+37% compared to 2000) and 7,779 red deer (+54%) have been harvested in 2004-
2005.
(Poster presentation.)
41
Interspecific competition between large herbivores: the fallow deer -
roe deer case.
P. Kjellander
Grimsรถ Wildlife Research Station, Dep of conservation biology, Swedish University
of Agricultural Sciences (SLU), Sweden
Integrated management of our Swedish four boreonemoral deer species, moose (Alces
alces), red deer (Cervus elaphus), fallow deer (Dama dama) and roe deer (Capreolus
capreolus), when occurring in the same area is highly requested but still missing. This
is unfortunate but most probably due to an almost complete lack of empirical studies
in Scandinavia on how guilds of ecologically similar species may coexist within the
same community. However, a prerequisite to create realistic management models
would be information not only on the relative influence of all the herbivores on its
resource but also measures of the interaction between the species themselves. This just
initiated project aim to investigate the type and strength of interactions (interference
and exploitive) which may occur among large herbivores in Sweden. This is not only
of high scientific value, but also of high importance in our ambition to learn more
about the possible effect of multispecies communities on forest production. Since the
strongest effects of competition among all the four sympatric Swedish deer species is
expected between the exotic fallow deer and naturally occuring roe deer, we will focus
our effort, in the short term, on that relationship. As an equally important by-product
of this project we will also be able for the first time to describe the general biology of
wild fallow deer (i.e. diet, seasonal habitat choice, home range size and seasonal
movements), the third economically most important hunted deer species in Sweden,
providing a basis for a sound future management of this species. This will be the first
time in Fennoscandia that the general biology of fallow deer will be studied in the
R 55
wild.
(Poster presentation.)
42
Current knowledge of the Central American red brocket deer
(Mazama temama Kerr, 1792) in Mexico.
J. Bello-Gutiรฉrrez
Divisiรณn Acadรฉmica de Ciencias Biolรณgicas. Universidad Juรกrez Autรณnoma de
Tabasco. Km. 0.5 carretera Villahermosa - Cรกrdenas. Entronque a Bosques de Saloya
CP 86039.Villahermosa, Tabasco, Mรฉxico
One of the deer species of which we have the least knowledge is the Central American
brocket deer (Mazama temama). The objective of this study is to review the
information produced regarding their biology and ecology. This deer is considered a
subspecies of M. americana, but is currently recognized as M. temama, having been
granted species status based on old karyotype data; segregating M. temama from M.
americana and including two subspecies; M. temama cerasina, present from Mexico
to Costa Rica, and M. temama reperticia, from Panama to Northern Colombia.
Information regarding population density is only available locally, varying from 0.09
deer/km
2
(including M. pandora) to 0.25 deer/km
2
in tropical forests, and 0.32 deer/
km
2
in cloud forests. The abundance index has been estimated using track counts,
ranging from 0.1 to > 1.8 track/km, in areas of sympatric with M. Pandora is higher
(> 1.8 track/km). In the Calakmul and Lacandona reserves, Red brocket is the second
most important source of food. Control hunting is practiced to reduce damage in beans
and corn fields. There have been seven births in the Chapultepec Zoo, with a 71.43%
survival rate, weight varied from 0.5 kg to 1.4 kg a few days after birth. A diet was
created for captive animals which represented 2 to 4 % of its body weight, and
included 16% crude protein, 2% lipids, 26% crude fiber and a Calcium-Phosphorus
relationship of 2.5:1. This deer is considered a representative of well-preserved forest
sites, however, it can be found in transformed sites. Further research is needed to
determine the degree of disturbance and isolation of the fragments that they inhabit
since habitat fragmentation, tourism, natural disasters and degradation by agricultural
activities are the main factors affecting potential habitats and distribution. This species
currently is not considered as threatened in the NOM-059-SEMARNAT-2000. It is
necessary to obtain more information on their habitat status, distribution, harvest
levels and abundance, as well as their importance to human communities in the areas
of subsistence and control hunting.
(Poster presentation.)
56 R
43
Energy requirement of captive grey brocket deer (Mazama
gouazoubira) determined by weight equilibrium and double-labeled
water.
A. Berndt
1
, M. Z. Moreira
2
, J. M. B. Duarte
3
, J. Barbosa
2
, and D. P. D. Lanna
2
1
Sรฃo Paulo Agribusiness Tecnology Agency - APTA, Andradina, Sรฃo Paulo, Brazil.
2
Superior Agriculture School ''Luiz de Queiroz'' - ESALQ/USP, Piracicaba, Brazil.
3
Veterinary Medicine and Zootecny School - FCAV/UNESP, Jaboticabal, Brazil.
There are limited data on energy requirements of brazilian cervids. Thus, it is difficult
to succeed in their management and reproduction in captivity. The objective of this
experiment was to study the nutritional requirements of the grey-brocket deer
(Mazama gouazoubira) in captivity. The determination of energy requirements for
maintenance used 8 gray-brocket deer of both sexes in captivity and was carried
through two methods: a)weight equilibrium and b)double-labeled water (2H218O).
The animals were dosed with double-labeled water (111.8 mg/kgBW for 2H2O and
163.1 mg/kgBW for H218O) and blood samples were collected with 3 days interval,
until 3 or 4 half lives of isotopes had occurred (reached limit of detection at
approximately 30 days after the dosage). The curves of isotopes disappearance as a
function of time were used to calculate the turnover of CO
2
and H
2
O. The results
obtained from the two methods were similar (111.4 and 112.0 kcal/kg.75.d) proving
the double-labeled water technique may be used in nutritional studies of cervids.
Information on doses (mg/kgBW) and maximum interval between injection and blood
collection (30days), allow the use of this methodology in future studies with free
ranging deer. This work determined the requirements of energy of grey-brocket deer
in captivity and validated the use of one indirect technique for use in free ranging
animals.
(Poster presentation.)
44
Modelling the influence of resources on the distribution and
aggregation of red deer hinds during the rut: implications for
mating system and management.
J. Pรฉrez-Gonzรกlez, A. M. Barbosa, and J. Carranza
Biology & Ethology, Universidad de Extremadura, Caceres, Spain.
Resource distribution is empirically known to affect the spatial distribution of female
red deer during the rut. We used geographical information systems and multivariate
R 57
analyses to quantify this relationship and assess which resources have the greatest
influence on red deer hind distribution in 35 populations in Mediterranean ecosystems.
We measured eight resource-related variables in each 50x50 m square, that included
distance to ponds, to dry streams, to green meadows, to food supplementation sites,
and to pasture areas, percentage of pasture cover, percentage of leguminous plants (in
pasture areas), and distance to croplands. For each population we built a stepwise
multiple regression model that resulted in a prediction of hind distribution based on
a selected significant subset of resources. Our results show that artificial food
supplementation and the presence of green meadows are the most important
determinants of hind distribution in these populations, although each resource may
have different importance in different populations. In addition, the spatial aggregation
of the most important resources for hind distribution was significantly related to the
spatial aggregation of hinds. This aggregation, apart from population traits such as sex
ratio and population size, is known to increase the degree of polygyny , therefore
reducing effective population size and tending to decrease genetic variability in
subsequent generations. Hence, the models relating female distribution to the
distribution of resources might be useful to predict how the management decisions
over particular resources are likely to affect relevant behavioural and evolutionary
parameters.
(Poster presentation.)
45
Red deer as a newcomer in Estonian fauna.
T. Randveer
1
and E. Niittee
2
1
Estonian University of Life Sciences, Institute of Forestry and Rural Engineering,
Tartu, Estonia
2
Ministry of Environment, Estonia
Cervus elaphus is a new species in Estonian fauna. The introduction of species started
in 1970-s when specimens from Voronez nature reserve, Latvia and Lithuania were
released on the islands of Saaremaa and Hiiumaa. Later the red deer reached South
Estonia moving there from Latvian territory. Appr. 1700. red deer was counted in
2006 in Estonia.
Forming the strategy how to treat towards the new species we must to find answers
to following questions:
1) How has the species adapted to our natural condition?
2) How is the sharing of habitats carried out with indigenous cervidae the roe deer and
moose and whether there are any signs of competition?
3) What is the influence of red deer on habitats, primarily upon forest renewal?
In the year 2000 a research project financed by the Centre of Environmental
Investments was started. The aim was the detailed study of the new species. In course
of it the following has been done:
58 R
The red deer was included in the list of monitored species, what means the analysis
of hunting statistics and examining of shot animals.
In two regions In Laasi (island Hiiumaa) and in Tรผndre (South Estonia) the spreading
of roe deer, moose and red deer in their winter habitats was estimated using the
method of counting the winter pellet groups. Additionally, during winters of
2000/2001 and 2001/2002 all it all 48 specimens of red deer were tracked and their
movings and all signs of vital activities were fixed and mapped by using the GPS. For
testing a hypothesis if the red deer`s moving are not random because of looking for
certain kinds of forage so we studied the distribution and density of tree species in
deer tracts and on randomly selected transects.
Our data show:
According to the data got by snow tracking and also by preliminary data got by rumen
content analysis the red deer consume economically important tree species in very
small amount. Most important food is grasses, forbs and dwarf-shrubs.
There is no evidence of existing acute competition for habitats between red deer and
other cervidae.
The condition of our red deer population is very good.
Resulting from the above and from the statement that the red deer is a very valuable
game species we can conclude that there is no reason to hinder the spreading of the
new species. In a wider perspective we consider desirable that red deer might become
the main game species in islands, which means also that the moose number must be
cut down by hunting. In Southern Estonia red deer probably remains species of
marginal importance.
(Poster presentation.)
46
Comparison of different weaning times of farmed Hungarian red
deer (Cervus elaphus hippelaphus) calves.
Z. Pados
1
, J. Szabรณ
1
, J. Nagy
1
, Sz. Nagy
2
, and Z. Zomborszky
1
1
University of Kaposvรกr, Faculty of Animal Sciences, H-7400 Kaposvรกr, Guba S. str.
40., Hungary
2
Research Institute for Animal Breeding and Nutrition, H-2053 Herceghalom,
Gesztenyรฉs str.1., Hungary
Research on the adaptation of deer farming technologies to Hungarian conditions
started in the 1980โs. Current practice in Hungary is to wean after the breeding season
mainly due to animal welfare concerns. However, due to economical reasons, there
is a growing interest in the application of earlier weaning techniques. To investigate
the effects of different weaning times under Hungarian conditions, we compared the
live weights of red deer (Cervus elaphus hippelaphus) calves weaned in August,
September or October in April in the following year. Data were collected in six
breeding centers (A-F) of the same company in Southwest-Hungary. Weaning times
R 59
were 08 August (AUG), 03 September (SEPT) and 29 October (OCT), 2004. Live
weight measurements were taken at weaning and in 21 April, 2005 (yearlings). A total
of 75 male and 77 female calves were measured. Data were analyzed with Statistica
for Windows 6.0. The effects of different weaning times as well as the different
breeding centers were analyzed with Kruskal-Wallis ANOVA followed by a post-hoc
Mann-Whitney U-test with Bonferroni corrections. Because sex had a significant
effect on April weights (median values for males and females were 74 and 64 kg,
resp., p<0.01), males and females were analyzed separately. There were no significant
differences between the April weights of different weaning times (AUG vs SEPT vs
OCT) in case of either sex (median values were 71 vs 75 vs 74 kgs, p=0.21 and 61 vs
64,5 vs 63 kgs, p=0.1 in males and females, resp.). ANOVA revealed a significant
difference among breeding centers in case of females (p=0.045) but not in males
(p=0.28). Post-hoc analysis revealed that the above mentioned difference was
significant only between centers B and D (p=0.002); all other differences were not
significant. The lack of significant differences between the live weights of calves with
early and late weaning dates in the following spring suggests that an early weaning
can be safely adapted.
(Poster presentation.)
47
A photographic guide for aging fallow deer Dama dama.
A. M. De Marinis, C. Gozzi, V. Marasco, and S. Toso
Istituto Nazionale per la Fauna Selvatica, Via Cร Fornacetta, 40064 Ozzano
dell'Emilia (BO), Italy
A photographic reference system for aging fallow deer Dama dama based on
permanent teeth eruption pattern and presence-absence of wear characters in jaws is
presented. Eruption pattern has been established by examining the jaws of 38 animals
(18 females and 20 males) of known age. The relationship between tooth wear and age
was studied counting cementum annuli in the root tips of I1 extracted from 35 jaws
of known age (22 females and 14 males). Incisors were demineralised, sectioned with
the use of a cryostat and stained with Ehrlich's haematoxylin. As the age assigned
reading cementum annuli was consistent with the known one of each jaw, we extended
the study to 94 animals (37 females and 57 males) of unknown age. Fallow deer were
shot (from November to May) in the Preserve of Castelporziano near Rome (Italy) as
part of the normal culling programme, between 2003 and 2005. As shown by the
ongoing researches in the Preserve all the fawns are born within a short 4-week period
between the end of May and the beginning of June. During this period some fawns
were ear-tagged with plastic tags; the tags were numbered and colour coded for the
year. The permanent teeth eruption sequence is as follows: I1, M1, M2, other incisors
and canines, P3 and other premolars, and M3. Full eruption is completed at 24-25
months of age. Studying permanent teeth eruption pattern we distinguished 4 age
60 R
classes: 5-6 months, 7-9 months, 10-12 months and 17-24 months, while studying
presence-absence of wear characters we used classes of 12 months. Males have higher
wear than females supporting the recent data on sex-specific life history tactics in
ungulates. The accuracy of the system was assessed through a blind test carried out
by three trained observers. The photographic reference system proposed can be easily
and quickly applied in the field and in routine data collections, keeping the time
required for aging each jaw to a minimum, but yielding reliable age determination.
However since times and sequences of eruption and tooth wear may differ between
areas, a calibration for each area is clearly recommended.
(Poster presentation.)
Diseases of deer
62 R
48
Recent advances in health and welfare of farmed deer in New
Zealand.
P. R. Wilson
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
Health and welfare of farmed deer in New Zealand are of internationally high repute,
but continuous improvement is essential to minimise clinical and sub-clinical losses
and to maintain market access for products. This paper summarises recent progress
with identifying, understanding and controlling diseases and initiatives underpinning
deer welfare. The national tuberculosis control programme reduced infected herds to
fewer than 50 in June 2005 a 32% reduction since June 2004, and has adopted new
rules and ancillary tests. Quality systems are employed to ensure optimum testing
standards. Johne's disease has emerged as the most significant current deer health
threat with implications for lesion differentiation from tuberculosis at slaughterhouses.
A substantial industry and government supported research and development project
is underway. Significant progress has been made in awareness and understanding of
this disease, and possible management strategies to reduce its impact. Leptospirosis
was recognised as a deer and human health concern resulting in significant research
to characterise the disease in deer and its epidemiology, and to investigate control
strategies. Vaccination reduces leptospiral shedding and kidney lesion and culture
rates. Preliminary observations have shown growth and reproductive responses in
some situations. Brucella ovis can cause high prevalence of disease in deer but a
research project characterised the disease and its epidemiology, identifying that self-
cure is common, and that an industry-wide control strategy was unwarranted since
there have been few recurrences. Recently, Cervid Herpesvirus-1 has been isolated
from deer eyes with keratoconjunctivitis, and from lesions in the vagina of deer. These
findings require further investigation. Recent evidence describes sub-optimum
efficacy of anthelmintics, and research shows the potential of forages for internal
parasite control. A national surveillance programme has confirmed no evidence of
Chronic Wasting Disease in New Zealand farmed deer. A number of veterinary and
laboratory reports of existing and novel clinical disease occurrences demonstrate
effective passive surveillance systems within the industry. Deer producers have
collaborated with industry stakeholders to develop a Deer Welfare Code that defines
minimum and preferred standards for the care of farmed deer, including health. This
code is currently undergoing regulatory approval, and thereafter will have legal
standing.
(Oral presentation.)
R 63
49
Health and production challenges facing intensive deer farming
industries.
P. R. Wilson
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
Deer farming industries in each country have specific consumer, economic,
management and health issues but many are common most. This paper briefly
addresses the key challenges likely to be faced by successful deer industries in the
future.
Consumer awareness of nutritional properties, production, welfare, product safety and
health will be a significant driving force for developments on-farm, but should be
regarded positively since solutions are generally compatible with improved production
and economics. Food safety concerns are universal, and will prompt a greater
requirement for assurance. Chemical usage and genetically modified substances,
organisms and possibly animals will be increasingly questioned. Natural alternatives
for disease, pest and parasite control, such as forages with health-promoting properties
and genetic selection for disease resistance as demonstrated for tuberculosis and
yersiniosis will increasingly be adopted. Farm management practices will be
increasingly scrutinised for animal welfare and environmental impact significance.
Failure to comply with community and consumer expectations for animal welfare risks
increased support for animal rights and liberation factions. Disease of real or perceived
significance possibly presents the greatest threat as demonstrated by consumer and
regulatory authority reaction to Chronic Wasting Disease. New diseases or cross-
species disease transmission, possibly with altered manifestation will continue to be
identified, and these must be researched to provide solutions to reduce risk to animals
and people. International trade will continue to be governed by phytosanitory
regulations but status may increasingly be at regional and possibly individual farm
level, requiring independent assurance. Economic trends dictate that continuous
production efficiency improvement will be necessary, often by increasing the scale of
operation, with its incumbent problems. Initial gains will be by conventional or novel
application of current technology such as forages and genetic selection for production
efficiency, and animal health plans including vaccine use to reduce wastage caused
by clinical and sub-clinical disease. Research will identify and refine systems, but on-
farm improvements will ultimately require more detailed multi-facet recording and
efficient information management systems. There will be an increased requirement for
professional advice and education to enhance the skill level of managers.
(Oral presentation.)
64 R
50
Chronic wasting disease in North America - A deer farmerโs
perspective.
C. Tedford
Tellico Plains, Tennessee 37385, USA
First observed in 1967 at a research facility in Fort Collins, Colorado; Chronic
Wasting Disease (CWD) did not affect deer farmers until discovered in herds of free
ranging and captive elk (wapiti) late in the 1990s. After almost a decade of extensive
sampling and testing, CWD has been identified in wild deer and elk herds in 13 states
and Canadian provinces and in captive deer and elk herds in 12 states and provinces.
Over 20 years of exhaustive study show no evidence that CWD poses any risk to
humans or domestic livestock. Despite this fact the necessary programs to monitor and
control the disease have placed an onerous burden on deer and elk farmers.
The regulatory burden combined with the misconceptions generated by the media with
their salacious appetite for the โkiller disease of the monthโ has dried up the pool of
potential deer farmers and driven many established farms out of business. An example
of this irresponsible journalism has been the implication in many articles and
broadcasts that humans could contract CWD or the related Creutzfeldt-Jakob by the
eating of venison.
Commercial herds numbering one thousand or more farmed deer and elk have all but
disappeared from the North American landscape. Over 10,000 small hobby farms
survive with the great majority consisting of less than 30 animals. Those of us who
remain with herd populations once considered potentially economically viable are
continuing with the hope that some of our allies in the research institutions of our state
and federal governments or private enterprise will discover an accurate and efficient
live animal test.
This test, along with the much needed epidemiological information that only
expensive research can provide, may remove much of the burden under which we now
must operate. Valuable research has already confirmed the mode of transmission of
the disease. In addition, two separate research projects, one conducted by our United
States Department of Agriculture Veterinary Services division and another by our
USDA Agricultural Research Service have indicated that it is possible that the Fallow
species of deer may possess a resistance to chronic wasting disease. Many other
projects are ongoing and promising.
(Oral presentation.)
R 65
51
Chronic wasting disease in Canadian wildlife: An expert opinion on
the epidemiology and risks to wild deer .
C. Maxwell
Canadian Wildlife Federation, 350 Michael Cowpland Drive, Kanata, ON K2M 2W1,
Canada
Introduction
The Canadian Wildlife Federation is one of Canadaโs oldest and largest wildlife
conservation organizations. For more than 40 years we have been promoting the
sustainable use of Canada's natural resources and the conservation of wildlife and its
habitat.
Along with Environment Canada, a department of Canada's federal government,
CWF co-sponsored the 2004 report entitled โChronic Wasting Disease in Canadian
Wildlife: An Expert Opinion on the Epidemiology and Risks to Wild Deerโ, more
commonly referred to as the Expert Panel on CWD. My presentation will focus on the
outcomes of this Expert Panel.
The Report:
โThe overall objective of the Panel is to provide an expert opinion on the best way
to research and manage CWD in wild deer populations in Canada. We hope that our
report will offer guidance to federal and provincial regulatory agencies in drafting
policies to contain or eradicate CWD in free ranging deer populations. A second but
equally important objective of the Panel is to provide a package of information to the
general public about risks associated with CWD based on data and experience gained
internationally in the last decade or so.โ
Specifically, the mandate of the Panel was to:
1) Improve the collective understanding of CWD in Canadian wildlife
2) Review risk factors and implications of CWD to wild cervid populations,
including future development of the disease throughout Canada
3) Provide an expert opinion on the potential risks of CWD to humans
4) Propose recommendations to manage impacts of CWD, focusing on surveillance
and monitoring programs, prevention, eradication, containment and human health
5) Encourage a national and international cooperative framework to assess risks
and manage CWD in wild deer populations
Panel Conclusions and Recommendations:
Recommendations, unanimous among members of the Panel, were divided into
4 parts, namely management of game farms, management of free-living cervids,
research needs and communications. Overall the Panel felt a sense of urgency in
taking actions to eradicate or at least contain CWD in Canadian wild deer populations.
66 R
CWF Concerns
CWFโs concerns stem from the unknown aspects of CWD. Of primary interest is
the range of susceptible species. We now know that moose are indeed naturally
susceptible to CWD, despite government assurances and best scientific guesses that
they are protected by the species barrier. We couldnโt agree more with the Panel that
research is needed into the susceptibility of caribou populations.
Environmental contamination, only confirmed as a reality within the last few
years, has us deeply concerned for the impacts of CWD on Canadaโs environment and
species.
The Panel states that a financial investment into CWD in the wild equal to that on
game farms is necessary to control or eradicate CWD. In an age of budget cutbacks,
we are worried that this will not occur.
The Future
CWF is encouraged by the Panelโs recommendations. We are hopeful that
adequate funds will be directed to all aspects of CWD containment, eradication and
research. CWFโs policy is clear that research on CWD should continue until the
disease is successfully eradicated from Canada.
The report Chronic Wasting Disease in Canadian Wildlife: An Expert Opinion
on the Epidemiology and Risks to Wild Deer can be downloaded or viewed at the
following website:
http://wildlife1.usask.ca/Publications/CWD%20Expert%20Report%20Final%20-
%2020040804.pdf .
(Oral presentation)
52
Epidemiological investigations of Johne's disease in deer.
J. C. Glossop
1
, P. R. Wilson
1
, C. Heuer
1
, and G. Nugent
2
1
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand.
2
Landcare Research, Lincoln, New Zealand
Johne's Disease (JD) has become an important disease in farmed deer herds in New
Zealand, causing losses of up to 20% of young growing deer on some farms, and
significant losses in older deer. A substantial research project is under way to quantify
the disease, and to identify, formulate, test and monitor practical and effective farm
management policies and practices to reduce its welfare impact upon animals and
economic impact upon farmers. Case control, crosssectional and cohort/longitudinal
study methodologies have been considered for this investigation. A nationwide
descriptive analysis and casecontrol study of 176 deer farms throughout New Zealand,
comparing farming practices between JD infected and clear herds was chosen as the
R 67
initial methodology, and preliminary results will be presented. Culture of six pooled
faecal samples from 10 mature deer each, estimated to have a herdbased sensitivity
of 68%, was chosen to define herd status, along with clinical and other evidence.
Further analysis of the sensitivity of this screening method is under way. All herds
with, and some without clinical evidence of JD were culture positive suggesting herd,
environmental, management and/or other factors may predispose to clinical disease.
Potential factors have been surveyed in detail for all participating farms and will be
subjected to multivariable statistical analysis to identify likely risk factors for
expression of disease. Key risk factors identified will be investigated using
intervention studies. Concurrently, wildlife on three properties with severe expression
of disease in deer have been hunted and necropsied, and tissues cultured for
Mycobacterium avium subsp paratuberculosis. The organism has been identified in
mesenteric lymph nodes or gut of 22% of wildlife animals sampled including rabbits,
hedgehogs, hares, possums, feral cats and ducks. The significance of these infections
will be investigated further. A national database to record lesion status from deer
slaughterhouses is being established to help define the present situation, to assist
slaughterhouse carcass inspection procedures, to monitor the effect of interventions
and adoption of new technology on infected farms in the future, and to monitor
progress toward control. A brief description will be presented.
(Oral presentation.)
53
Johne's disease in farmed deer in New Zealand.
C. G. Mackintosh
1
, J. F. T. Griffin
2
, and G. W. de Lisle
3
1
AgResearch Invermay, Mosgiel, New Zealand
2
Department of Microbiology and Immunology, University of Otago, Dunedin, New
Zealand
3
AgResearch Wallaceville, Upper Hutt, New Zealand
Johneโs disease, caused by M. avium subsp. paratuberculosis, has emerged as an
important problem in farmed red deer (Cervus elaphus) in New Zealand and other
countries. It causes financial losses due to poor growth rates and death in young deer,
it increases the rate of non-specific reactions to the skin test for tuberculosis (Tb) and
interferes with normal meat inspection procedures in deer slaughter premises by
causing Tb-like lesions in mesenteric lymph nodes. Infection has been confirmed by
culture on ~10% of New Zealand deer farms. Unlike the disease in sheep and cattle,
Johneโs disease in farmed red deer commonly occurs in young animals, 8-15 months
old. Affected animals lose weight and condition over a relatively short period and
develop diarrhoea terminally. Typical gross signs at necropsy include enlarged
mesenteric lymph nodes, which may contain caseous lesions, thickened intestinal
mesenteries, often with cord-like lymphatic ducts and mild to moderate thickening of
the small intestine. Histopathological examination reveals varying degrees of
68 R
granulomatous enteritis, which may be multi- or paucibacillary. Clinical disease is
most likely to result from heavy oral challenge in young animals. Although the ovine
strain of M. paratuberculosis has been isolated from deer, it appears that most disease
problems are associated with the bovine strain. Diagnosis depends on combinations
of gross signs, serological tests, histopathology, culture and PCR. A commercial IgG1
ELISA has been shown to be relatively sensitive and has been useful for screening
affected herds and reducing clinical disease in seriously affected herds. Pooled faecal
culture offers a cost-effective means of detecting infected herds. Tissue culture
remains the gold standard for differentiating between bovine tuberculosis, avian
tuberculosis and paratuberculosis. Improved, cost-effective methods are required for
the control of Johneโs disease in farmed deer. Control of Johneโs disease currently
relies on reducing the exposure of young animals to heavy challenge by culling
clinically affected animals, optimising nutrition and minimising stress. Screening
hinds with the IgG1 ELISA is useful for culling subclinically infected deer, because
pregnant hinds infected with M. paratuberculosis are very likely to pass infection onto
the unborn foetus. In future, vaccination may assist in reducing losses if problems of
sensitisation to tuberculin testing can be overcome.
(Oral presentation.)
54
Insights into the pathogenesis of Johneโs disease in red deer (Cervus
elaphus).
C. G. Mackintosh
1
, J. Thompson
1
, J. F. T. Griffin
2
, and G. W. de Lisle
3
1
AgResearch Invermay, Mosgiel, New Zealand
2
Department of Microbiology and Immunology, University of Otago, Dunedin, New
Zealand
3
AgResearch Wallaceville, Upper Hutt, New Zealand
Research over the last 5 years has resulted in a clearer understanding of the
pathogenesis and epidemiology of M. avium subsp. paratuberculosis infections in
farmed red deer (Cervus elaphus) in New Zealand. Using an experimental challenge
model, we showed that oral challenge of young red deer with a high dose (10
9
cfu) of
bovine strain M. avium subsp. paratuberculosis resulted in clinical disease in
approximately a third of animals, whereas medium (10
7
cfu) and low doses (10
3
cfu)
did not cause clinical Johneโs disease but resulted in dose-related degrees of
subclinical disease. The ovine strain appears to be less virulent for red deer than the
bovine strain of M. paratuberculosis. Pregnant red hinds are very likely to pass
infection onto their offspring either in utero or via infected milk. Necropsy and culture
of 9 clinically affected hinds in the last trimester of pregnancy showed that the
foetuses of 8/9 hinds were infected. Similarly, 14/18 subclinically infected hinds in the
second half of pregnancy had infected foetuses and M. paratuberculosis was isolated
from the mammary tissue or lymph node of 13/16 of these subclinically infected
R 69
hinds. A challenge trial showed that vaccination with a killed M. paratuberculosis
mineral oil adjuvanted vaccine significantly reduced the severity of paratuberculosis
lesions. However, the vaccine significantly interfered with ante mortem skin testing
for bovine tuberculosis. Over 90% of vaccinated and challenged animals gave a false-
positive reaction to the single intradermal skin test with bovine tuberculin, but all the
animals gave avian responses to a comparative skin test conducted 13 months after
vaccination.
(Oral presentation.)
55
The efficacy of oral and pour-on ivermectin and pour-on moxidectin
in farmed red deer.
S. O. Hoskin
1
, W. E. Pomroy
1
, P. R. Wilson
1
, M. Ondris
1
, and P. Mason
2
1
Institute of Veterinary, Animal and Biomedical Sciences, Massey University, New
Zealand
2
Mason Consulting, New Zealand
The efficacies of ivermectin (IVM); oral (IVMo) and pour-on (IVMp), and pour-on
moxidectin (MOX) were determined using forty 4-month-old weaner red and wapiti,
red deer naturally infected with both lungworm and gastrointestinal (GI) parasites.
Deer were maintained on ryegrass pasture in mid-autumn and randomly allocated
based on liveweight, sex, genotype, faecal egg and larval count, to 4 treatment groups
(n=10) which were control (no anthelmintic), oral IVM (200ยตg/kg), pour-on IVM
(500ยตg/kg) or pour-on MOX (500ยตg/kg). Faecal egg (FEC) and larval (FLC) counts
were taken at -3, 0, +7 and +10 days post treatment. At Day +10 post-treatment deer
were necropsied for total worm counts. In the control group the geometric mean (GM)
burdens of Dicytocaulus spp., Teladorsagia-type spp., Oesophagostomum spp. were
4552, 244 and 196 respectively. No Haemonchus, few Trichostrongylus axei and no
small intestinal nematodes were found. Arithmetic mean worm burdens were generally
similar to GM burdens and reductions in AM burdens are summarised here.
Anthelmintic efficacy against lungworm ranged from 99.98-100% despite high
numbers of larval lungworm in control deer. Reduction in FLC at +10d was 100% for
all anthelmintics. At Day +10 small numbers of lungworm were recovered from lungs
of 50% of animals treated with IVMo and 20% of animals treated with MOX. Efficacy
against Oesophagostomum was 100% for all anthelmintics. Efficacy against total
Teladorsagia-type spp. from the abomasum were: pour-on MOX 94.1%; IVMo
31.1%; IVMp 68.5%. Nematodes were recovered from the abomasums of all deer
treated with both IVM formulations and 60% of deer treated with MOX. Efficacies
against Teladorsagia leptospicularis, S. kolchida, Spiculopteragia spiculoptera and
S. assymmetrica varied between anthelmintics with MOX totally effective against
Spiculopteragia spp. FEC reductions (corrected for change in control FEC) were
MOX and IVM pour-on 100% and IVMo 56.3%. The sub-optimum efficacy against
70 R
Teladorsagia-type nematodes with IVMp, the formulation licensed for use in deer,
requires further investigation.
(Oral presentation.)
56
An internatinal review of Leptospirosis in wild and farmed deer.
M. A. Ayanegui-Alcรฉrreca
1
, P. R. Wilson
1
, C. Heuer
1
, J. M. Collins-Emerson
1
, C. G.
Mackintosh
2
, A. C. Midwinter
1
, and F. Castillo-Alcala
1
1
Institute of Veterinary, Animal, and Biomedical Sciences, Massey University, PB
11222, Palmerston North, New Zealand
2
AgResearch Invermay, PB 50034, Mosgiel, New Zealand
Leptospira infections in wild and farmed deer occur worldwide. This paper reviews
technical literature on leptospiorisis in deer first available from 1957 to 2005 including
66 reports from 16 countries. Forty-five reports (68%) were from wild deer including
four that were leptospira negative, sixteen were from farmed deer including nine from
New Zealand, three describing experimental infection, and two from zoos. Most
(29.5%) were from red (Cervus elaphus), with 18% from fallow (Dama dama), 17%
from white-tailed (Odocoileus virginianus) and 10% from roe deer (Capreolus
capreolus). Most reports (73%) described serology only, with evidence for 19 serovars
including Pomona (26%), Hardjobovis (15%), Copenhageni (15%), Grippotyphosa
(11%) Ballum (6%). Culture was reported only for serovars Hardjobovis, Pomona,
Copenhageni and Roumanica. Disease, culture and gross and/or histopathological
lesions were described in 18 (27%) reports associated with serovars Pomona,
Hardjobovis, Copenhageni, Roumanica and Ballum. Proof of disease causation
requiring culture, is described only for Pomona, Copenhageni and Roumanica. Most
reports of disease were from farmed deer in New Zealand, describing haemolysis,
jaundice, renal lesions, haemoglobinuria and sudden death in individuals or outbreaks
attributed to Pomona. Mixing of young stock from several sources appears a
significant risk factor. Subclinical kidney lesions are associated with Pomona and
Hardjobovis. Slaughterhouse workers are at greatest risk of contracting disease from
deer. Vaccination produces serological responses, but effectiveness in protecting from
disease, and reduction of shedding in urine, remains unreported. Evidence suggests
that various deer species can act as a reservoir (maintenance) host for Hardjobovis, an
accidental individual and alternative maintenance population host for Pomona and
possibly accidental hosts for serovars Roumanica and Copenhageni. Insufficient
evidence exists for establishment of the host:organism relationship for other serovars.
More robust knowledge of leptospiral occurrence in many countries, epidemiology of
infections, and the effectiveness of vaccines and vaccination regimes is needed to
assist farmed deer industries to develop leptospirosis management strategies.
(Oral presentation.)
R 71
57
Epidemiology of Leptospiral infections with Serovars Hardjobovis,
Pomona and Copenhageni in farmed red deer (Cervus elaphus) in
New Zealand.
M. A. Ayanegui-Alcรฉrreca
1
, P. R. Wilson
1
, C. G. Mackintosh
2
, J. M. Collins-
Emerson
1
, C. Heuer
1
, A. C. Midwinter
1
, and F. Castillo-Alcala
1
1
Institute of Veterinary, Animal, and Biomedical Sciences, Massey University, PB
11222, Palmerston North, New Zealand.
2
AgResearch Invermay, PB 50034, Mosgiel, New Zelaand.
This paper reports two longitudinal studies of the epidemiology of leptospirosis in
farmed deer herds in New Zealand and a national prevalence survey. A serum bank
of 3-monthly samples from each of 10 rising 1-year-old, 12-24-month-old and adult
deer on 16 farms during 1992-3 was used for a retrospective sero-epidemiological
analysis. All 16 farms and 51.8% of individuals were seropositive for Hardjobovis
with seroprevalence lowest in 4-month-old deer and highest in deer >1-year-of-age.
Four farms (25%) and 24.6% of individuals were seropositive to Pomona with the
pattern within and between farms resembling epidemic infection. Two farms (13%)
and 10.5% of individuals were seropositive for Copenhageni but evidence suggested
cross-reactivity to serovar Pomona. In 2003-4 a second study involved approximately
6-weekly sampling over one or two production cycles for serology, urine culture and
urine dark field microscopy (DFM) from predominantly young deer from one farm
infected with serovar Hardjobovis, and two with dual Hardjobovis and Pomona
infections. Kidneys were collected from deer at slaughter. Hardjobovis seroprevalence
ranged from 7.5-57.5% and titres ranged from 0-3072. Pomona seroprevalence ranged
15-97.5% with titres up to 12,228. There were mild kidney lesions but no disease
associated with Hardjobovis, but evidence of disease and severe kidney lesions in 68%
of deer with Pomona. Up to 60% of urine samples were DFM positive with shedding
persistence exceeding nine months. Infection patterns suggest that deer are
maintenance hosts for Hardjobovis and accidental individual and possibly
maintenance population hosts for Pomona. Twelve to 20 blood samples/herd from 9-
to 30-month-old unvaccinated red and red x wapiti deer farms throughout NZ (n=110;
80 sampled on-farm, 30 sampled at slaughterhouses) were collected March 2003 to
February 2005 for leptospira serology. Overall, 82% of herds were infected. At the
herd and individual animal levels, respectively, seroprevalence for Hardjobovis was
77.7% and 60.8%, and for Pomona 20% and 8.4%. Within-herd seroprevalence for
both serovars ranged from 0 to 100%. Concurrent Hardjobovis and Pomona infections
were observed in 16.4% of herds and 6.6% of individuals. There were no regional
differences in seroprevalence.
(Oral presentation.)
72 R
58
Anthelmintic use and internal parasite control in farmed deer in
New Zealand.
F. Castillo-Alcala, P. R. Wilson, W. E. Pomroy, and S. O. Hoskin
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
A questionnaire was posted to 500 deer farmers in New Zealand in November 2004
to gather information about parasite control yielding 227 replies (45.4%) with 198
suitable for analysis. Ninety four percent of respondents used anthelmintics at least
once in the 12-month period defined with 53% treating all deer classes and 22%
treating young deer only. Seventy four percent based anthelmintic dose on weight of
the heaviest animal, with 36% using a weigh scale. Deer up to 1-year-of-age (n=175
farms) were treated 1-13 times (mean 3.2) starting from January to November. Mean
treatment interval ranged from 41- 46 days. Yearling and adult hinds and stags were
treated at least once (range 1-7) on 55-64% of farms, depending on animal class.
Moxidectin was the most commonly used anthelmintic (46-58%, depending on animal
class), followed by abamectin, eprinomectin, oxfendazole, ivermectin, albendazole,
levamisole and doramectin with selection based mainly on perceived efficacy. Weight
gain and body condition were the most common means for monitoring parasitism in
young and older deer, respectively. Few respondents used faecal egg and/or larvae
counts. Coughing and/or scouring were diagnosed as parasitism in young deer on 13-
14% of farms, and deaths associated with lung and gastro-intestinal worms were
recorded on 5% and 3% of farms, respectively. Veterinary diagnosis occurred in 23%
of events. Production losses and/or death of yearling and/or adult deer due to
parasitism were reported by 27% of respondents. When planning anthelmintic
treatment programmes, 63% of respondents followed advice from veterinarians. Thirty
four percent always placed deer on clean or spelled pastures after treatment, while
32% did that often. Fifteen percent had incorporated forages and/or herbs with
presumed anthelmintic properties into their parasite control programme. Forty four
percent were very confident of a return on investment when using anthelmintic on
their deer. Respondents stated that their knowledge of the life cycle of the major
parasites of deer was very good (8%), reasonably good (61%), poor (28%) and nil
(3%). Results demonstrate variability of parasite control practices for farmed deer in
New Zealand and suggest that improvements could enhance productivity and
profitability.
(Oral presentation.)
R 73
59
Subdural occurrence of Elaphostrongylus cervi and Setaria cervi in
red deer of West Hungary.
L. Sugรกr, Sz. Kovรกcs, and A. Kovรกcs
Kaposvรกri University, Faculty of Animal Science, Hungary
Veszprรฉm University, Georgikon Faculty of Agronomy, Hungary
Prior final settlement, young specimens of Elaphostrongylus cervi and Setaria cervi
nematodes visit the subdural space of the cranial cavity in red deer (Cervus elaphus
L). To evaluate the frequency, intensity, seasonality and possible pathological
consequences of this subdural activity 196 red deer from six West Hungarian
populations were examined in the period of March 1998 and March 2006. Deer heads
were bisected sagittally and the surfaces of brain and dura mater observed for worms
and disorders. E.cervi worms were found in all deer populations examined with values
of prevalence from 18.2 to 46.7%. The highest prevalence was observed in the
population living inthe area of Danube flood-plain called Gemenc. The mean intensity
(worm burden) varied between 1.4-2.3 worm in all populations, with an individual
maximum of 6 in two calves. In regard of the host age, calves were the most affected:
36.3% prevalence and 1.79 mean intensity. Then the decreasing values indicate the
evolvement of the solid immunity. The presence of S servi was usually infrequent
3.16% overall prev, with a surprisingly high value, 40.7% in a sample (n = 24) in
January 2000 in Zala county. Both worm species occurred between October and
February only, except one calf in June 2001, with three E. cervi specimens. The gross
pathological alterations were restricted to the inner surface of dura mater; yellow
infiltration, colorless translucent, pearl-like application. Because clinical symptoms
were never observed, our opinion is that E. cervi seems to be a harmless parasite of
red deer.
(Oral presentation.)
60
Disease problems in Mongolian reindeer.
J. C. Haigh, M. G. Keay, V. Gerwing, J. Erdenbaatar, and M. Nansalmaa
Western College of Veterinary Medicine, University of Saskatchewan; Canada
During two (2004, 2005) one-month summer field trips into the taiga regions of north-
western Mongolia where the Tsaatan people herd their reindeer we took whole blood
samples from 147 and 97 animals respectively. The samples were either preserved in
EDTA or allowed to clot. Serum was harvested from the clotted samples after they had
been allowed to stand for 30 min at ambient temperature. As the region is at least a
74 R
weeks travel from laboratory facilities we carried out initial screening for Brucellosis
using Brewer card test kits on site within 24 hours of sample collection. Serum and
blood was stored in sterile vials and maintained in coolers using river water or packed
snow. In 2005 parallel Rose Bengal tests were carried out on sera using a Mongolian
antigen at the State Central Veterinary Laboratory in Ulaanbaatar. There was a reactor
rate in excess of 23% in both years. A limited number of whole blood samples were
subjected to DNA extraction and primers for Brucella spp. were used to detect the
presence of this organism. Because a Brucella Strain 19 live vaccine vaccination
program was carried out under government mandate in the mid-1980s we are
endeavoring to distinguish between this organism, which can cause a self-sustaining
infection in reindeer, and other possible strains, including B. suis Biovar 4. Blood
smears were made and stained in the field using Diff-Quik stain and stored for
transportation to the laboratory. There they were examined under oil immersion at
100X. An unidentified hemoparasite was seen in approximately 10% of slides. We
collected milk samples from 27 reindeer and one sample of a sero-sanguineous
material aspirated under sterile conditions from a swollen carpal joint. DNA and
bacterial culture was conducted on these samples. Udder lesions with the appearance
of cow pox were seen on 17 reindeer at two camps in 2004. We carried out further
clinical examinations on 14 and 12 individual reindeer in the two years. Of these, 10
were for lameness and /or swollen joints. Six cases of saddle sores were seen at three
camps.
(Oral presentation.)
61
Histopathology of fluorotic coronal dentine of roe deer (Capreolus
capreolus) and red deer (Cervus elaphus) teeth.
H. Richter, A. Richards, and H. Kierdorf
Department of Biology, University of Hildesheim, Marienburger Platz 22, 31141
Hildesheim, Germany (HR, HK)
Royal Dental College, University of Aarhus, 8000 Aarhus C, Denmark (AR)
Intake of excessive amounts of fluoride during dental development causes
pathological dental changes, known as dental fluorosis. Dental fluorosis is used as a
biomarker of elevated fluoride exposure of wild deer. The present contribution
summarises pathological changes of dentine structure and mineralization in fluorotic
teeth of roe and red deer from fluoride polluted regions in Central Europe. Mandibles
were selected based on their dental lesion index of fluorosis and compared to non-
fluorotic specimens. From each specimen ground sections of P4, M1 and M3 were
analysed using light microscopy and microradiography.
In fluorotic dentine, long period incremental markings were accentuated in the form
of hypomineralised bands. In severely affected specimens, single or multiple broad
bands of interglobular dentine were present, occurring preferentially in the dentine
R 75
below the infundibulum and near the enamel-cementum junction. In some cases, bands
of interglobular dentine could be followed through the entire coronal dentine. In places
it could be demonstrated that hypomineralized long period incremental markings were
continuous with bands of interglobular dentine. Both phenomena are regarded as
denoting different degrees of impairment of the process of matrix mineralization,
occurring along the dentine mineralization front. It was possible to match
pathologically enhanced striae of Retzius in the enamel with accentuated long period
incremental markings in the dentine. So-called Owens contour lines were found to be
associated with interglobular dentine. It is hypothesized that in the affected specimens,
fluoride exposure constitutes an additional stress factor responsible for an increased
susceptibility of the odontoblasts, leading to a deviation from their normal course.
Different degrees of pathological dentinal changes occurred among different teeth of
one individual, with the M1 showing the least severe alterations. This situation can
hypothetically be related to the developmental sequence of the permanent dentition.
Since dentine formation continues throughout the life of an individual, the analysis of
fluorotic changes in dentine allows an assessment of the fluoride exposure of an
animal throughout life.
(Oral presentation.)
62
Mineral composition and requirements for growth of farmed red
deer in New Zealand.
F. Castillo-Alcala
1
, P. R. Wilson
1
, and N. D. Grace
2
1
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
2
AgResearch Grasslands, Palmerston North, New Zealand
This paper presents the first substantial data on the concentrations, quantities and
requirements for growth of Ca, P, Mg, Na, K, Cu, Mn, Fe and Zn in farmed red deer
in New Zealand. Six normal neonatal, four 12-month and four 20-month-old deer
averaging 9.6, 68.25 and 94.7 kg, respectively, were dissected, and organs (brain,
lungs, heart, spleen, liver, kidney, pancreas), offal, digestive tract (stomach, small and
large intestines), muscle, bone, skin and blood weighed and sub-sampled for mineral
determinations carried out by inductively coupled emission spectrometry. Data are
presented on the concentrations and amounts of each mineral in each organ and tissue,
and estimates of total content and concentration against empty body weight. Expressed
as a percent of total mineral in the empty body weight bone contained 99% of Ca, 88%
of P, 59% of Na and 59% of Mg, muscle contained 82% of K, 38 % of Fe and 62%
of Zn. The organs contained less than 6% of the minerals with the exception of Fe, Mn
and Cu. The lungs, spleen and liver had a high concentration of Fe and the digestive
tract a high concentration of Mn. Each kilogram live weight gain was associated with
14.5 g Ca, 10.7 g P, 0.54 g Mg, 1.1 g Na, 2.4 g K, 1.56 mg Cu, 0.63 mg Mn, 42.7 mg
76 R
Fe and 26.6 mg Zn. The distribution of Cu between the liver, muscle and bone was 77,
8 and 3% for the newborn and 24, 59 and 2% for older deer. Data can be used to
estimate mineral requirements, using a factorial model, adopting assumptions from
other species for absorption and endogenous losses pending deer-specific data. Thus
the Cu requirement of a deer growing at 180g/d can be estimated as 0.18 kg x 1.56 mg
(mg/kg live weight gain) = 0.28 mg, plus endogenous loss (7g (estimated loss/kg) x
68 kg) = 0.48 mg, divided by the estimated absorption coefficient (0.05) and daily DM
intake (2.2 kg/day). The requirement is therefore estimated to be at least 7 mg/kg DM.
Confirmation requires controlled feeding studies.
(Oral presentation.)
63
Recent advances in understanding therapy with Copper Oxide Wire
Particles in New Zealand Farmed deer.
P. R. Wilson
1
, F. Castillo-Alcala
1
, and N. D. Grace
2
1
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand.
2
AgResearch Grasslands, Palmerston North, New Zealand
This paper summarises investigations into the distribution of Copper Oxide Wire
Particles (COWP) and effect of molybdenum on their efficacy in red deer. In study 1,
10g COWP were given to 10 of 20 deer grazing pastures containing low, medium or
high Mo concentrations for 92 days from late October. Treatment maintained adequate
mean serum Cu concentration in deer grazing low and medium Mo pastures (< 5
mg/kg DM) for >92 days but <50 days in deer on high Mo pastures (8-12 Mo/kg DM)
(p=0.003). Peak and overall mean liver Cu concentrations in deer on high Mo pasture
were lower than in those grazing low and medium Mo pastures (p<0.001). Thus
elevated dietary Mo reduces absorption and therefore storage of dietary Cu, requiring
higher supplementation frequency. Study 2 investigated gastro-intestinal distribution
and faecal excretion of particles of a novel copper oxide wire particle (COWP)
product (CUE Bullet). Twenty 18-month-old red deer hinds were treated orally with
a bolus of 10g COWP and four slaughtered on days 1, 5, 15, 30 and 60 after treatment.
The gastro-intestinal tract was secured between compartments and contents rinsed
until sedimentation of particles occurred. Sediment was oven dried and copper
particles were separated and weighed. Faeces were collected continuously from four
additional animals in metabolism cages for four days after treatment, and sub-sampled
for daily particle recovery estimates. Copper oxide wire particles were found in all
compartments of the gastro-intestinal tract caudal to the oesophagus for at least 15
days and in the rumen/reticulum and abomasum for at least 60 days. After 24 hours,
a mean of 6.21g of COWP were recovered, declining to 2.6, 0.46, 0.52 and 0.15 after
5, 15, 30 and 60 days, respectively. The highest recovery rate at every sampling day
was from the rumen/reticulum (55-97%). A mean of 0.09g of COWP (range 0 0.24g)
R 77
was recovered from faeces during the first 24 hours. Over four days, a mean of 0.94
g (range 0.03 2.61g) was recovered from faeces. These studies have added to
knowledge of factors affecting the efficacy of COWP, and their dynamics in the
gastrointestinal tract.
(Oral presentation.)
64
Nasopharyngeal bot fly, Oestridae larvae in red deer in Hungary.
L. Sugรกr, Sz. Kovรกcs, and A. Kovรกcs
Kaposvรกri University, Faculty of Animal Science, Hungary
Veszprรฉm University, Georgikon Faculty of Agronomy, Hungary
Bot fly larvae (Oestridae, Diptera) develop for about six month period in the nasal
cavity and later in the pharynx of the host. In red deer (Cervus elaphus L.) two species
occur in general Cephenemyia auribarbis Meigen 1824 and Pharyngomyia picta
Meigen, 1824. The joint occurrence of the two species was examined in a total of 138
red deer of West Hungary between March 1998 and March 2006. Larvae were
detected and collected after the midsagittal cutting of the head. The species
identification, prevalence and number of larvae (intensity), and the variation due to
host age and season were analyzed by statistical methods. The overall prevalence was
92,7%, with 22 median intensity if the two bot fly occurrence data were combined.
However the prevalence of C. auribarbis was only 46.9 % in contrast to 90.6% for P.
picta due to the elongated development of the later species. The highest values were
found in the calves and yearlings: 97,3% and 100% prevalence 17,0 and 6,0 median
intensity respectively (what are in conflict with the results experienced in Spain
previously). The evaluation of the seasonal occurrence of larvae shows clearly the
differences in the seasonal development characteristics of the two bot fly species. C.
auribarbis larvae reach their maturation tightly together (February-March), therefore
seemingly earlier, than the great mass of the P. picta larvae (March-June). It was often
difficult if not impossible to count the total number of larvae in the host, especially in
regard of P. picta, because most of the first stage larvae staged in the lower airways
(trachea and bronchi). In regard to the bot fly larvae we did not see signs of obvious
acquired immunity in contrast of the situation experienced with the warble fly larvae
(Hypoderma spp.). The general pathogenic or weakening effect of the bot fly larvae
could not be proved too.
(Oral presentation.)
78 R
65
ITS2 sequences of Dictyocaulus lungworms from red and fallow
deer in Hungary: molecular evidence for a new genotype.
Z. รcs, L. Sugรกr, and Z. Pรฉnzes
Faculty of Animal Science, University of Kaposvรกr, Guba S. Str. 40. PO. Box 16.
Kaposvรกr H-7400, Hungary
Infection caused by lungworms (Dictyocaulus spp.) in deer is a well known problem
for wildlife managers and deer farmers. Lungworms are considered to be the most
important parasites in deer (Cervidae). There are evidences for exist a few
Dictyocaulus species in deer in Europe (D. eckerti, D. viviparus, D. capreolus, D. sp.
from fallow deer), but it is hard to distinguish them accurately on the basis of their
morphological features. In our study, sequencing of the second internal transcribed
spacer (ITS2) of nuclear ribosomal DNA (rDNA) was applied to the genetic
characterisation of Dictyocaulus collected in different Hungarian red deer (Cervus
elaphus) and fallow deer (Dama dama) populations. The ITS2 of Dictyocaulus from
fallow deer and part of red deer congeners with previously published sequences of D.
eckerti. However, lungworms from red deer derived from central Transdanubia
differed in sequence from D. eckerti and D. viviparus by 38% and 30% respectively.
The ITS2 sequence data indicates that it is as genetically different as are the
mentioned species of Dictyocaulus. It was thought until that date, the specific
Dictyocaulus parasite of red deer is D. eckerti, although D. viviparus occured in some
cases as well. These data show evidence for existing of a significantly differed
lungworm genotype living in red deer. It is indicated there is real need for a large-
scale molecular systematic study of Dictyocaulus specimens from red deer (and other
cervids) for epidemiological studies.
(Poster presentation.)
66
Fascioloidosis of red deer and its therapy in ''Szigetkรถz'' region in
the North-West of Hungary (1998-2005).
B. Egri and E. Giczi
University of West Hungary, Department of Animal Health, Mosonmagyarรณvรกr,
Hungary
The giant liver fluke (Fascioloides magna Bassi, 1875)was detected for the first time
in Hungarian red deer shot in 1994 and became extensive up to date.The parasite can
be found in the region extending from the Czech Republic, through Slovakia and from
2000 into Austria. The mean prevalence of fascioloidosis among red deer in Hungary
R 79
between 1997-1998 was 72.0-60.7%. In the period of 1998-2005, at necropsy of 459
deer livers (using Egri's method), the number of flukes per host ranged from 1 to 138.
The first attempts for game treating was performed until 2000, using the Rafendazole-
premix(with rafoxanide)
-medicated feed in feeding places and feeding boxes. The efficiency was low (mean
prevalence:51%). The efficacy of later used other preparations ( SBH-
Exwormer(SBH)(with triclabendazole + levamizole) or Tribex (Chanelle) (with
triclabendazole)) - feeding method was the same - were evaluated in the treatment of
naturally acquired fluke-infections in red deer. The efficiency of preparations in
different years was different. Using the QP 2.0 method of the quantitative
parasitology, the typical extreme values of eight years were the following: Confidence
limits of prevalence:9.55-78.5%,mean prevalence values:44.19%, and the confidence
limits of median intensity were:1-77. Index of Discrepancy was:0.641-0.896.
Differences between the data on prevalence-level of the eight years were statistically
significant, because ''P''(chi square test for comparing prevalence) was 0.016.
(Poster presentation.)
67
Coprological monitoring of Trematodes in free-ranging red deer
population at eastern Croatia.
A. Slavica, T. Florijanฤiฤ, Z. Janicki, D. Konjeviฤ, K. Severin, R. Beck and K. Pintur
Veterinary Faculty, University of Zagreb, Croatia
During the four year period (2001-2004) a coprological monitoring was applied on
free-ranging red deer population at eastern Croatia, with purpose to detect presence
of trematodes developmental stages in faeces. Dropping samples of deer were
collected monthly according to different areas (hunting grounds). All samples were
analyzed in laboratory by standard flotation and sedimentation method and at the same
time eggs per gram (EPG) determination were done. Over four years in nine hunting
grounds total number of 2017 samples was collected. Eggs of three trematode species
were detected, i.e. large American liver fluke (Fascioloides magna), liver fluke
(Fasciola hepatica) and ruminal trematode (Paramphistomum cervi). In one hunting
ground (''Podunavlje-Podravlje'' - referral number XIV/9) maximum percentage of
positive samples was detected for P. cervi (78,3 %) with high EPG value (max. = 556,
0 = 48,5), F. magna have lower positive percentage (52,5 %) with lower EPG value
(max. = 300, 0 = 41,5) and F. hepatica have just 9,3 % of positive samples with max.
of 55 EPG (0 = 17,2). In five hunting grounds (''Radinje'', ''Koha-Kozarac'', ''Haljevo'',
''Munjoroลก'', ''Podravlje'') we determined no presence of F. magna eggs, while P. cervi
eggs were detected in 45 % (mean value) of samples from all five hunting grounds
with EPG = 31. Presence of F. hepatica eggs were detected in 11,9 % of samples and
mean value of EPG for all five hunting ground was low (<5). In three hunting grounds
(''ล arkanj-Vrblje'', ''Spaฤva-South'', ''Spaฤva-North'') we found low ratio of F. magna
80 R
eggs (12,3 %) with EPG = 12,5, while P. cervi eggs were found in 53 % of samples
with EPG = 33,5 and F. hepatica eggs were detected in 7,9 % of samples with EPG
= 5. Correlation between antitrematod treatment and EPG values for all found
trematodes is also given in this paper.
(Poster presentation.)
68
Sub-clinical parasitism, weaning date, growth of deer fawns and
reproductive performance of hinds .
J. M. Mwendwa, M. L. W. J. Broekhuijse, S. O. Hoskin, W. E. Pomroy, and P. R.
Wilson
Institute of Veterinary Animals and Biomedical Sciences, Massey University,
Palmerston North, New Zealand
A recent survey indicated March was the most common month for pre-rut weaning of
farmed deer in New Zealand (31%), with 5% weaned in February (n=119 farms). Both
weaning date (March vs. June) and date of first anthelmintic treatment potentially
influence liveweight gain of young deer. This study investigated the impact of pre-rut
weaning date on parasitism and liveweight gain (LWG) of red deer fawns and hinds,
and conception date and rate of hinds by untrasound scanning. Seventy-six deer fawns
were randomly allocated in a 2x2 factorial design, involving weaning date (February
17 or March 17) and two treatments with moxidectin anthelmintic at a six-week
interval (Jan 14 and Feb 25) or no anthelmintic treatment. LWG of fawns was
monitored at 2-4 week intervals from Jan 12 to Mar 31. Sixty-four mixed-age hinds
were used to investigate the effect of pre-rut weaning date on internal parasitism and
conception date. Deer grazed permanent perennial ryegrass-based pasture together
until weaning. Fawns weaned in March had a higher LWG to March 31 than those
weaned in February (P<0.0001). A significant weaning by anthelmintic treatment
interaction was found (P<0.02), with LWG being higher in treated fawns weaned in
March (P<0.017) but not February (P>0.10). Faecal larval counts (FLC) in treated
fawns were zero, but faecal egg counts (FLC) in treated fawns averaged 136 epg
(range 0-600). In hinds, FLC averaged 5 lpg (range 0 โ 122) and FEC averaged 26
(range 0- 200) with no significant relationship between weaning date and either FLC
or FEC. No effect of weaning date was shown on conception rate or date. This study
has shown that pre-rut weaning date and sub-clinical parasitism during summer and
early autumn can influence LWG in young farmed deer.
(Poster presentation.)
R 81
69
Investigation of the sanitary status of red deer (Cervus elaphus)
culled in the Italian Alps between 2001 and 2005.
E. Andreoli
1
, I. Bertoletti
2
, A. Bianchi
2
, E. Heinzl
1
, E. Scanziani
3
, and S. Mattiello
1
1
Istituto di Zootecnica, Faculty of Veterinary Medicine, University of Milan, Via
Celoria 10, 20133 Milan, Italy
2
IZSLER, Sezione Diagnostica di Sondrio, Via Bormio 30, 23100 Sondrio, Italy
3
DIPAV, Faculty of Veterinary Medicine, Universitร degli Studi di Milano, Via
Celoria 10, 20133 Milan, Italy
The sanitary monitoring of wild populations is desirable to improve knowledge about
the epidemiology of those diseases present, to provide guidelines for a preventive
medicine programme, to protect the health of people who work with those animals and
to preserve the health balance between the different species present in the study area,
especially in alpine environments, where spatial overlap may occur between wild and
domestic ruminants in summer ranges. The health status of wild red deer of Fontana
Valley (where red deer and domestic ungulates coexist during the summer grazing
period), in the province of Sondrio (Italy), has been monitored since 2001. Gross
examination was carried out on 143 animals culled from 2001 to 2005, during the
hunting season (September-November), and body conditions were scored from 1
(poor) to 3 (good). Blood sera were obtained from blood collected from the heart and
were submitted to serological analysis, while tissue samples from organs including
heart, lungs, kidney and diaphragm were submitted to histopathological analysis.
Arcview 3.2 was used to provide topographic maps to monitor the spread of the two
most common diseases in the study area, namely parasitic bronchopneumonia and
Sarcosporidiosis, to evaluate the incidental influence of the presence of the domestic
ruminants and to highlight the areas of greatest risk of transmission. Epidemiological
index of prevalence was calculated for each disease. The most important diseases
found in the study population are Leptospirosis (Leptospira serovars
Australis/Bratislava, Grippotyphosa, Pomona, Seyroe/Hardjo,
Icterohemorragiae/Copenhageni ), Bovine Respiratory Syncytial Virus, parasitic
bronchopneumonia and Sarcosporidiosis. The prevalence is generally low, except for
bronchopneumonia (74.1%) and Sarcosporidiosis (63.3%). Brucella abortus/melitensis
was recorded by the rose bengal plate agglutination test. However, complement
fixation test could not confirm these results, because of an anticomplement activity of
these sera. The general conditions of the study population were good, in spite of the
relatively high prevalence of some diseases. This suggests that red deer in the Fontana
Valley are coping well with the present situation, but topographic maps show that they
may represent a risk for the transmission of diseases to domestic livestock, and
highlights the importance of the sanitary monitoring of wild populations.
(Poster presentation.)
82 R
70
General comparison of taxonomic characters distinguishing two
closely related species of deer lice - Solenopotes burmeisteri and S.
capreoli (Phthiraptera, Linognathidae).
V. Bรกdr, P. ล tindl, and J. Preisler
Dept. of Biology, University of Hradec Krรกlovรฉ, Rokitanskรฉho 62, 500 03 Czech
Republic
The history of description of two deer lice is discussed. Adults of Solenopotes
burmeisteri (Fahrenholz, 1919) and S. capreoli Freund, 1935 are redescribed and the
male of S. capreoli described and illustrated for the first time. The species are
compared on the basis of major taxonomic characters: morphology of head, thoracic
sternal plate, male and female genital area, abdominal chaetotaxy and body
measurements. Both the lice have been noted as very rare species, S. capreoli have
been observed only in three European countries; this may be due to inadequate
collecting technique. Dissolving deer hides in 5% potassium hydroxide solution seems
to be the only accurate method of assessment of prevalence and infestation
intensity.Both species of lice are localized predominantly on the hosts head, around
the eyes and on the cheeks.
(Poster presentation.)
Genetics and Evolution
84 R
71
Landscape features affect gene flow of Scottish Highland red deer
(Cervus elaphus).
S. Perez-Espona, J. McLeod, F. J. Perez-Barberia, C. G. I. Jiggins, and J. Pemberton
Institute of Evolutionary Biology - The University of Edinburgh, UK
Despite Scotland holding the largest population of red deer (Cervus elaphus) in
Europe, population genetics studies have been mainly undertaken in the Island of
Rum. For red deer inhabiting mainland Scotland, in particular the Highlands where a
more semi/natural population exists, studies assessing effective dispersal (gene flow)
and population genetic structure are lacking. In this study we assessed the population
genetic structure of Scottish Highland red deer by analysing 695 individuals for 21
microsatellite markers. Additionally, we also assessed the influence of several natural
and man-made landscape features on red deer gene flow in the Scottish Highlands by
following a landscape genetics approach and using GIS techniques. Despite the
relatively small scale of the study area, significant population structure was found
using F-statistics (FST = 0.019) and the Bayesian clustering analysis implemented in
the program STRUCTURE (K = 4). Population genetic structure analyses, isolation
by distance analyses and the Monmonierโs algorithm implemented in the program
BARRIER indicated the Great Glen valley as the major gene flow barrier in the study
area. Landscape features were shown to significantly affect red deer gene flow as they
explained a greater proportion of the genetic variation than geographical distance
between populations. Sea lochs were found to be the strongest red deer gene flow
barriers in our study area, followed by roads, mountain slopes and forests. Inland lochs
and rivers were identified as red deer gene flow corridors.
(Oral presentation.)
72
Sex biased dispersal in an expanding red deer population.
H. Haanes, K. H. Rรธed, and O. Rosef
Norwegian School of Veterinary Science, Dep of Basic Sciences and Aquatic
Medicine, PO-8146 Dep, N-0033 Oslo, Norway
Telemark University College, Dep of Environmental and Health Studies, N-3800 Bรธ
in Telemark, Norway.
The demography and distribution of species vary through time and space and greatly
affect levels of genetic variation and population structure. We have investigated the
genetic variation and differentiation of 14 microsatellites in the Norwegian red deer
subspecies (Cervus elaphus atlanticus), which has expanded demographically and
R 85
spatially the last hundred years after a major population decline from 300 to 200 years
ago. Significant F
st
-values from 0.021 to 0.197 between localities sampled across the
present distribution indicated moderate to strong genetic structure and a Mantel test
showed highly significant isolation by distance. Similar levels of genetic variation and
recent bottlenecks indicated that genetic drift during the population decline formed the
present genetic structure. Natural barriers to gene flow consituted by steep fiords and
high mountains has maintained isolation by distance during population expansion.
This was supported by Bayesian assignment tests that showed a 99.9% probability of
five Norwegian subpopulations, four of which were concurrent with the distribution
of four of main locations during the population decline. The assignment tests further
revealed that first-generation dispersal was heavily sex-biased with twice to six times
as many males as females. Division of the data-set into five clusters according to the
assignments test, F
st
values among males were significantly lower than among
females, suggesting that female matrilinear groups have kept together during the
spatial expansion. Female matrilineal grouping in red deer thus even seems common
during a spatial population expansion.
(Oral presentation.)
74
A molecular phylogeny of the evolutionary radiation of New World
deer (Odocoileinae, Cervidae): Implications for biogeography and
the evolution of antlers.
S. M. Carr
1
, E. D. Richards
1
, H. D. Marshall
1
, and J. M. Smith-Flueck
2
1
Genetics, Evolution, and Molecular Systematics Laboratory, Department of Biology,
Memorial University of Newfoundland, St. Johnโs NL A1B3X9, Canada
2
National University of Comahue, Bariloche, Argentina
Molecular phylogenetic relationships among 11 genera of telemetacarpalian โNew
Worldโ Cervidae were investigated with the mitochondrial cytochrome b gene and the
more slowly evolving 12S rRNA gene. Telemetacarpalian deer, which comprise โNew
Worldโ Odocoileinae and antlerless Old World Hydropotes, can be divided into three
monophyletic tribes: Capreolini (Capreolus and Hydropotes), Alceini (Alces only),
and Odocoileini (holarctic Rangifer and endemic nearctic and neotropical deer).
Antlers evolved only once: Hydropotes shows a secondary loss of antlers, and does
not represent the plesiomorphic state for cervids. The data challenge conventional
hypotheses about the evolution of New World cervids. Odocoileus, the typical nearctic
deer genus, extends into Central America, and the occurrence of O. virginianus in
South America north of the Amazon basin has been taken to suggest that all South
American deer evolved recently from O. virginianus. Molecular analysis shows
instead that the endemic large-bodied South American genera, comprising Marsh Deer
(Blastocerus), Pampas Deer (Ozotoceros), and Huemul & Taruca (Hippocamelus
spp.), are part of a monophyletic clade that diverged from the North American clade
at the time of the reconnection of the Panamanian Isthmus 3.5 MYBP. Small-bodied
86 R
Pudus (Pudu) are the sister group to this clade. In contrast, the small-bodied Central
and South American Brocket Deer (Mazama spp.) are more closely related to
Odocoileus, and represent a secondary invasion of the neotropics. Relationships within
Mazama are complex. Reduced ''spike'' antlers in Brocket Deer and Pudus appear to
be parallel consequences of allometry for reduced size, rather than an indication of
close relationship.
(Oral presentation.)
75
Genetic distinctiveness of isolated and threaten Tsaatan reindeer
herds in Mongolia.
K. H. Rรธed, J. C. Haigh, V. Gerwing, and M. Keay
Norwegian School of Veterinary Science, Oslo, Norway
The Tsaatan and their reindeer have historically nomads throughout the Sayan border
region, covering a taiga home range that stretched far north of todays Siberian-
Mongolian border. The community historically interacted with neighbouring ethnic
groups, swapping deer as needed. In the 1930-50s, the Tsaatan group fled south to
Mongolia after which the border was firmly closed and all contact with other herds
and ethnic groups stopped. The herd on the Mongolian side was probably around 2-
3000 reindeer according to a 1977 estimate. In the following decades the size of the
herd was further reduced to probably only a few hundred individuals for thereafter to
increased and during the last 15 years the herd size has been around 6-700 animals.
The population is split between the two taiga settled areas called east and west taiga,
between which virtually no interaction occurs, and these are further split into different
family herds. Here we present data on the genetic variation of the east and west
Tsaatan reindeer herds by analyses of allelic variation of 14 microsatellite DNA
markers together with sequence variation of the control region of mitochondrial DNA.
The Tsaatan reindeer herds are genetically very distinct compared to other reindeer
across Eurasia. The herds are characterized by significantly reduced amount of genetic
variation probably attributable to bottleneck processes with subsequent effects of
inbreeding and genetic drift.
(Oral presentation.)
R 87
76
Conservation genetics of Argentinean pampas deer populations.
S. Gonzรกlez
1
, M. Cosse
1
, V. Raimondi
2
, M. L. Merino
3
, B. Galvan
4
, and J. E.
Maldonado
5
1
Departamento de Genรฉtica-IIBCE Unidad Asociada Facultad de Ciencias UdelaR
Av. Italia 3318 Montevideo 11600- Uruguay
2
Cรกtedra de Genรฉtica. Departamento de Ciencias NaturalesFacultad de Humanidades
y CienciasCiudad Universitaria. Paraje el Pozo s / n. Universidad Nacional del
Litoral. Argentina.
3
Div. Zool. Vertebrados. Museo de La Plata/ CICPBA . Paseo del Bosque s/n La Plata
B1900FWA Buenos Aires- Argentina.
4
Universidad Nac. de La Pampa, 6300 Santa Rosa- Argentina.
5
Genetics Program, National Zoological Park/National Museum of Natural History,
Smithsonian Institution,Washington, D.C. 20008- USA
Populations of the endangered Pampas deer have been dramatically reduced
particularly in the southern part of its range where only four small and highly isolated
populations remain in Buenos Aires (Bahรญa Samborombรณn), Corrientes, Santa Fe and
San Luis provinces of Argentina. In order to deduce genetic units for conservation and
to better understand the effect of habitat fragmentation on gene flow and genetic
variation, we performed a molecular genetic study of three populations of Pampas deer
from Argentina and compared them to samples obtained from other South American
localities. In total, we examined 100 individuals using two mitochondrial DNA
markers, the control region and the cytochrome b gene. Analysis of the control region
revealed that pampas deer had high levels of polymorphism reflecting large historic
population sizes of millions of individuals in contrast with the low numbers observed
today. The three Argentinean populations analyzed showed high levels of
polymorphism and did not share haplotypes. In spite of having unique haplotypes, the
populations from San Luis and Bahรญa Samborombรณm did not show significant levels
of genetic differentiation suggesting that they belong to the same genetic unit. The
cytochrome b marker revealed a star like phylogenetic network with a central common
haplotype present in all the populations from Brazil, Argentina and Uruguay. The
direct ancestor of the Pampas deer probably first appeared in the Ensenadan
stages/ages during the Pliocene about 2 million years ago. The phylogenetic pattern
suggests a range expansion in the pampas deer in the Pleistocene. The current
demographic decline is so recent that genetic variation has not yet been depleted. The
molecular genetic results provide a strong mandate for habitat restoration and to plan
alternative management strategies to preserve the levels of genetic variation and
recover the historic patterns of abundance.
(Oral presentation.)
88 R
77
Genetic characterisation of roe deer (Capreolus capreolus)
population of Parma Apennines.
C. S. Soffiantini
1
, G. M. Pisani
1
, M. Malacarne
1
, G. Gandolfi
2
, A. Sabbioni
1
, and J.
Tagliavini
2
1
Dipartimento Produzioni Animali, Biotecnologie Veterinarie, Qualitร e Sicurezza
degli Alimenti - University of Parma, Italy
2
Dipartimento di Biologia Evolutiva e Funzionale - University of Parma, Italy
According to historical reports, the roe deer (Capreolus capreolus) population of
Parma Apennines should be constituted of allochthonous subjects (coming from
Central Europe and from Slovenia) introduced for hunting purposes during the first
half of 1800. As a result of the improvement of conservative hunting management and
the decrease of intesive agricolture activities the consistency of roe deer stock
dimension markedly increased from 1950s in Parma. For the same reasons a continuos
habitat arose between allochthonous population of Parma and authochtonous
population of Southern Tuscany, with possible repecussion on expected genetic
structure of roe deer populations. The aim of this research was to study the genetic
structure of roe deer population of Parma Apennines by mitochondrial DNA
variability. In this concern, 40 mtDNA D-Loop (about 600 bp) taken from roe deer
tissue samples collected during the hunting season 2005-2006, were sequenced. We
have observed two main mitochondrial aplotypes that clusterised, respectively, with
central Italy mtDNA aplotypes (>70%) and Central Europe/East Alps (<30%). Nuclear
characterisation based on microsayellites variability to confirm these genetic
distribution is in progress.
(Poster presentation.)
78
Aplotypic characterization of roe deer by asymmetric PCR and
SSCP analysis.
J. Tagliavini, S. Casagrande, M. Malacarne, and P. Mariani
Dept. Biol. Evol.& Funz. and Dept. PABVQSA. Univ. of Parma, Italy
Mitochondrial D-Loop sequences of 40 roe deer (Capreolus capreolus), representative
of population of Parma Apennines (I), exhibit two main aplotypes that shows high
similarity respectively with sequences of Central-Italy (subspecies C. capreolus
italicus) and of East-Alps and Central-Europe (subspecies C. c. capreolus). The
alignment of D-Loop sequences have revealed four conserved nucleotidic mutations
which discriminate the two aplotypes, localized in a 200 bp length region. A pair of
R 89
primers were designed to amplify this variable region with whom we have performed
asymmetric PCR. The PCR samples, mainly single strand DNA synthesized by
forward primer, were mixed with two volumes of loading buffer (95% formamide, 20
mM EDTA, bromophenol blue and xylene cyanol) and loaded on an 8%
nondenaturing polyacrylamide gel, containing 5% glicerol and 1xTBE buffer.
Electrophoresis was carried out at 20C, at 10 V/cm for 6 hr, and then the gel was
stained with ethidium bromide. Clear different electrophoretic patterns with SSCP
(Single Strand Conformation Polymorphism) analysis were detected for different
aplotypes. The method is fairly rapid and inexpensive, and could be used to screening
large samples before carry out more expensive analysis.
(Poster presentation.)
79
Phylogeography of Iberian red deer populations and their
relationships with main European red deer lineages.
J. L. Fernรกndez-Garcรญa
1
, J. G. Martรญnez
2
, L. Castillo
3
, and J. Carranza
3
1
Dpto. de Zootecnia, Facultad de Veterinaria, Univ. Extremadura, Cรกceres SPAIN.
2
Dpto. de Biologรญa Animal y Ecologรญa, Facultad de Ciencias, Univ. de Granada,
18071 Granada, SPAIN
3
Cรกtedra de Biologรญa y Etologรญa, Facultad de Veterinaria, Univ. Extremadura,
Cรกceres SPAIN.
The Iberian subspecies Cervus elaphus hispanicus is morphologically distinct (lighter
and smaller) from the other western subspecies of red deer, but few samples have been
so far used to study its variability at mtDNA level and its relationships with other
western red deer populations. Here we use the first hipervariable segment of
mitochondrial D-loop (control region), which has proved to be a very informative
marker for defining subspecies boundaries. We screened 229 samples in total, of
which 195 were from four main Iberian areas (Extremadura in the west, Castilla-La
Mancha in central Spain, Andalusia in the south and Pirineans in the north), together
with 34 samples from other European areas (Britain, Norway, Germany and Balkans).
Additionally, some GenBank sequences were incorporated for haplotype matching and
phylogenetic analyses. A total of 53 different haplotypes were obtained, which
revealed two distant genetic lineages within Spain: one in the west and another in
central Spain. Some of our samples were obtained from trophies hunted during the
first half of XX century, presumably before most recent translocations took place.
Haplotypes from these samples provided support to the geographic distribution of
current native populations, and allowed us to detect some cases of exceptionally rare
haplotypes sampled in current populations that matched with foreing, distant
populations, hence presumably resulting after recent reintroductions. We suggest that
large areas in Iberia where the two genetic linages still persist should be preserved
from genetic admixture by preventing translocations.
90 R
(Poster presentation.)
80
The artificial occurrence of the fallow deer, Dama dama dama (L.,
1758), on the island of Rhodes (Dodecanese, Greece): insight from
mtDNA analysis.
M. Masseti
1
, A. Cavallaro
2
, E. Pecchioli
3
, and C. Vernesi
3
1
Dipartimento di Biologia Animale e Genetica ''Leo Pardi'', Laboratori di
Antropologia ed Etnologia, Universitร di Firenze. Via del Proconsolo, 12, 50122
Firenze, Italy
2
Via G. Barellai, 18. 50137 Firenze, Italy
3
CEA, Centro di Ecologia Alpina, Viote del Monte Bondone, 38040 Trento, Italy
The aim of the present work is to investigate the origin of the fallow deer of the island
of Rhodes Dama dama dama (L., 1758) (Dodecanese, Greece), by molecular means,
and survey this population for phenotypic variability. Our results show that these deer
have homogeneous phenotypic patterns. All specimens fell within the common colour
coat variety typical of the wild form. The Rhodian deer appear to be rather small,
especially if compared with specimens from central and northern Europe. We then
sequenced the HVR-I of 13 deer from Rhodes and compared these sequences with
other 31 samples obtained from different European and Anatolian populations of
fallow deer. Out of 44 sequences, 23 haploypes were found. When compared to the
Turkish and Italian population, the population of Rhodes revealed lower values of
within population genetic diversity. The fallow deer from Rhodes are characterized
by an 80bp mtDNA insertion not found elsewhere. As a consequence all the deer from
Rhodes form a tight cluster, distinct from all other fallow deer populations. This
uniqueness makes the conservation and management of the Rhodian population
particularly urgent.
(Poster presentation.)
R 91
81
Comparative anatomy of three Asian ruminant animals.
J. Kimura
1
and K. Fukuta
2
1
College of Bioresource Sciences, Nihon University, Japan
2
Graduate School of Bioagricultural Science, Nagoya University, Japan
The suborder Ruminantia can be divided into two infraorders, Tragulina and Pecora.
Tragulidae occupies a basal position with respect to all other ruminant families. The
considerable phylogenetical differences between Tragulidae and the other, more
recently evolved, ruminants have been established by means of DNA sequence
analysis. In this study, the comparison of the anatomical characteristics of the salivary
glands and the placenta of three ruminants in Asia were attempted.
Lesser mouse deer (Tragulus javanicus) were obtained from a breeding farm in East
Malaysia. Reeveโs Muntjac (Muntiacus reevesi) which had been introduced from
Taiwan were hunted in Chiba prefecture, Japan. Sika deer (Cervus nippon) were
hunted in Kanagawa prefecture, Japan. Their salivary glands and placentas were
dissected out and their histological structure was observed. The weight of the parotid
glands relative to the body weight was calculated. Among the three species, the lesser
mouse deer has the heaviest parotid glands (2.5g/kg). Reeveโs Muntjac was 2.0g/kg,
while the Sika deer was 0.8g/kg. There is a significant difference between mouse deer
and sika deer and also between muntjac and sika deer. The data on the lesser mouse
deer is the largest compared with other ruminants investigated by Kay (1987). The
function of the salivary glands in the lesser mouse deer must be most significant as it
is a concentrated selector in the ruminant animals. It has been speculated that the
significance of the salivary function changed through adaptation and evolution with
the selection of food intake. The gross structure of the placenta in the mouse deer was
diffuse and thus noticeably different from that of the muntjac and sika deer which are
polycotyledonary. Histologically, however, the placenta of Tragulidae appears to be
epitheliochorial and therefore similar to other ruminants. Numerous trophoblastic
binucleated cells, characteristically present in all other ruminants, were observed.
These results suggest that the placenta of Tragulidae is a transitional type between
diffuse epitheliochorial and polycotyledonary synepitheliochorial categories.
(Poster presentation.)
92 R
82
Characterization of the growth curve of red deer (Cervus elaphus
scoticus) in a herd in Central Mexico.
A. C. Delgadillo, R. Lรณpez, H. H. Montaldo, J. M. Berruecos, A. Luna, and G. C.
Vรกsquez
Department of Genetics and Biostatistics, Faculty of Veterinary Medicine and Animal
Husbandry, National Autonomous University of Mexico. and National Center for
Research in Physiology and Animal Improvement, INIFAP, Mexico.
Introduction
Red deer (Cervus elaphus) has changed from being considered as a cynegetic
species to an alternative species for meat production (Van den Berg y Garrick, 1997),
therefore description of the growth pattern in a defined environment and production
system, allows assessment of the importance of some factors that affect the productive
efficiency. These curves and their estimation from different functions (Behr et al.,
2001; Huisman et al., 2002), have been widely discussed in the literature on domestic
animals (Behr et al., 2001; Kaps et al., 2000; in cattle; Bathaei and Pascal, 1998 in
sheep; Huisman et al., 2002 in pigs; Mignon-Grasteau et al., 2000 in poultry; Lรณpez
et al., 2000 in fish; and Landete-Castillejos et al., 2001 in Iberian red deer). The
objective of this study was to compare four different models in order to characterize
growth curve in red deer within a herd in the state of Queretaro, Mexico.
Material and methods
The information corresponded to the weight records of 240 deer born between
2000 and 2002 taken every two months from birth to two years of age. Four models
were used with unadjusted weights within each sex in order to obtain the growth
curves, which were then compared using the determination coefficient (R
2
), square
mean of the error (SME) and Mallowsโ Cp statistic (Cp).
Results and discussion
Least square means of weights at birth, 95, 240, 380 and 660 days are shown in
Table 1, where it can be seen that males exceeded females by 8%, 14%, 14%, 17%
and 39% respectively (P < 0.05). Table 2 shows the equations and comparison criteria
taking into consideration R
2
, SME and Cp. We observed that the equations obtained
from the non-linear model. Within the non-linear models that estimate the parameters
with a biological perspective, the one that best fit the data was Brodyโs model. The
parameter of weight at maturity varied from 63 to 69 Kg in females and 105 to 130 Kg
in males in the different models with biological interpretation.
R 93
Table 1. Least square means grouped by sex for weight at birth (WB), weight at 95
days (WW), weight at 240 days (W8M), weight at 380 days (WY) and weight at 660
days (W2Y)
Trait (kg) N FEMALES N mALES
WB 112 8.82 + 0.13ยช 126 9.56 + 0.12
b
WW (95 days) 66 33.71 ยฑ 0.65ยช 81 38.47 ยฑ 0.59
b
W8M (240 days) 260 44.91 ยฑ 0.46ยช 301 51.01 ยฑ 0.43
b
WY (380 days) 136 55.41 ยฑ 0.86ยช 182 65.10 ยฑ 0.78
b
W2Y (660 days) 32 65.56 ยฑ 2.54ยช 177 91.40 ยฑ 1.08
b
a,b, means with different superscript in each row are statistically different, p<0.05
n= number of records
Table 2. Growth curves Characterization in red deer
SME R
2
Cp
Brody (exponential)
F y= 69.0075(1-0.8552 exp (-0.00392 x )) 42.45 0.978 76.15
M y= 130.5(1-0.8959 exp (-0.00158 x )) 95.69 0.974 281
Von Bertalanffy (sigmoid)
F y= 64.7309 ( 1 - 0.4414 exp ( -0.00589 x ))
3
45.9 0.976 130.78
M y= 110.2 ( 1 - 0.4727 exp ( -0.00296 x ))
3
104 0.972 379.24
Richards (sigmoid)
F y= 63.6237 ( 1 + 0.0660 exp (-0.00676 x ))
24.6297
47.47 0.975 157.83
M y= 106.0 ( 1 + 0.0602 exp (-0.00357 x ))
29.3409
107.5 0.971 423.64
Gompertz (sigmoid)
F y= 63.5100 exp (-1.6732 exp (-0.00687 x)) 47.69 0.975 159.27
M y= 105.6 exp (-1.8116 exp (-0.00364 x)) 107.9 0.971 426.36
Females(F), Males (M); means square error (SME), coefficients determination (R
2
)
and Mallowsโ Cp statistic; x= age in days
The weight at maturity varied from 63 to 69 Kg in females and from 105 to 130
94 R
Kg in males. A model by Brody standardized by the number of observations indicates
that the weight at maturity in females is 60 Kg and males is 132 Kg. been daily gain
if 855 and 899 g.The results suggest that Brodyโs model is the one most adequate for
characterization of the growth curve of red deer in this study, with the comparison
criteria that are being used.
References
Bathaei, S.S., Pascal, L.L. (1998) Genetic and phenotypic aspects of the growth curve
characteristics in Mehraban Iranian fat-tailed sheep. Small. Rumin. Res. 29: 261-269.
Behr de, V., Hornick, J.L., Cabaraux, J.F., Alvarez, A., Istasse, L. (2001) Growth
patterns of Belgian Blue replacement heifers and growing males In commercial farms.
Livest. Prod. Sci. 71:121-130.
Huisman A. E., Veerkamp R. F. and Van Arendonk J. A. M. (2002) Genetic
parameters for various random regression models to describe the weight data of pigs.
J. Anim. Sci. 80:575-582.
Kaps, M., Herring, W.O., Lamberson, W.R. (2000) Genetic and environmental
parameters for traits derived from the Brody growth curve and their relationship with
weaning weight in Angus cattle. J. Anim. Sci. 78:1436-1442.
Landete-Castillejos, T., Garcรญa, A., Gallego, L. (2001) Calf growth in captive Iberian
red deer (Cervus elaphus hispanicus): effects of birth and hind milk production and
composition. J. Anim. Sci. 79:1085-1092.
Lรณpez, S., France, J., Gerrits, W.J.J., Dhanoa, M.S., Humphries, D.J., Dijkstra, J.
(2000) A generalized Michaelis-Menten equation for the analysis of growth. J. Anim.
Sci. 78:1816-1828.
Mignon-Grasteau, S., Piles, M., Varona, L., de Rochambeau, H., Poivey, J.P., Blasco,
A., Beaumont, C. (2000) Genetic analysis of growth curve parameters for male and
female chickens resulting from selection on shape of growth curve. J. Anim. Sci.
78:2515-2524.
Van den Berg, G.H.J., Garrick, D.J. (1997) Inheritance of adult velvet anther weights
and live weights in farmed red deer. Livest. Prod. Sci. 49:287-295.
(Poster presentation.)
R 95
83
Mitochondrial DNA variability and polymorphism of ISSR-PCR
markers in the reindeer population of Eastern Siberia.
N. V. Kol, O. E. Lazebny, and I. A. Zakharov
Vavilov Institute of General Genetics, Russian Academy of Sciences, Russia
Mitochondrial DNA variability was analyzed in one of the southernmost isolated
populations of domestic reindeers (Rangifer tarandus L.) in the Republic of Tuva (in
south Eastern Siberia, Russia). DNA isolated from 29 skins of reindeers from Tuva
was used to analyze mitochondrial DNA polymorphism. A 470 bp fragment of the
control region of the D-loop of mitochondrial DNA was amplified and sequenced.
After alignment, a 418 bp sequence was taken for subsequent computer analysis.
Conservative regions were revealed in the sequence under study: at the beginning B
between base pairs 1 and 150 and at the end B between base pairs 355 and 418. In
variable regions transitions were mainly revealed B 23 T-C substitutions, 34 C-T
substitutions and 4 A-G substitutions. When comparing all nucleotide sequences under
study the number of variable sites was found to constitute 3,5% of the total number
of nucleotides. Eight haplotypes were found among 29 studied tuvinian samples.
Intrapopulational nucleotide diversity in the studied population was determined,
p=0,00859 ยฑ 0,00234. For comparison with our material sequences from GeneBank
belonging to 3 known mtDNA haplogroups. Suppose that haplogroup III has the east-
Siberian origin. Our data agree with their inference. The evolutionary relations
between the sequences D-loop of mtDNA were analyzed using the package of
computer programs. The results of statistical analysis of suggest the existence of at
least eight nucleotide positions characterized by a high level of homoplasia B from six
to three parallel mutations per position. The estimates evolutionary age of individual
groupings of mtDNA types in the tuvinian reindeers. The age of these three
evolutionary lines exceeds a presumable time of domestication of reindeers.
Moreover, 62 animals of the tuvinian reindeers were investigated with the help of
ISSR-PCR method. It has been revealed 71 fragments of various lengths. Also indexes
of average paired similarity and value of average heterozygosity are counted. Values
of average heterozygosity for first marker is 0,7307, and for another - 0,7488. This
values of the studied population comprised with the results of the microsatellite DNA
investigations of the reindeer populations. Thus, reindeers of the south of Eastern
Siberia keep high enough value of heterozygosity than island populations of reindeers
of Spitsbergen and Greenland, and also caribou Canada. The tuvinian population was
reduced significantly last years, so it is necessary to watch the genetic variety
preservation providing steady maintenance of a population.
(Poster presentation.)
96 R
84
A new conservation genetic union from Pampas deer (Ozotoceros
bezoarticus) in Southern Brazil.
F. G. Braga, S. Gonzรกlez, and J. E. Maldonado
Federal University of Paranรก, Rua Saldanha Marinho 1923, Curitiba - 80.730-180,
Brazil
The pampas deer is an endangered species in the state of Paranรก, southern Brazil. In
the past, this species was widespread in this state and inhabited the open grasslands
and the Brazilian cerrado. However, in recent times, destruction of their habitat has
caused populations to decline in numbers and the few populations that remain are
fragmented and isolated. We performed a genetic study to analyze the genetic
variability of a population from Piraรญ do Sul District, which is the largest known
population of pampas deer in southern Brazil with an estimated population size of 71
individuals. Samples from tissues were extracted from dead animals found in the field
and from skulls kept by ranchers and poachers in this area. Tissue samples (50 mg or
100 l) were transferred to 1.7 ml eppendorf tubes containing 95% ethanol. Procedures
for DNA extraction was done and no-template polymerase chain reaction (PCR)
controls were used in each amplification. Universal primers Thr-L15910 and DL-
H16498 were used in PCR reactions to amplify a 601 bp fragment. In addition, a 486
bp fragment of cytochrome b gene was amplified for using primers L14724 and
H15149. Purified PCR products were sequenced using the ABI Big Dye ready reaction
kit and ran on an automated sequencer ABI 377. Sequences were aligned using Clustal
X. Of the 51 samples that were extracted, 11 samples were successfully amplified and
sequenced for the D-loop region and 8 for the cytochrome b gene region. The 11 D-
loop sequences resulted in 4 different haplotypes and the 8 cytochrome b sequences
resulted in 2 different haplotypes. We compared these haplotypes with previously
published D-loop sequences and from unpublished cytochrome b sequences from
Brazil, Argentina, Paraguay and Uruguay. The results of our analysis revealed that all
4 D-loop and the 2 cytochrome b haplotypes were different from all the mitochondrial
haplotypes reported. Furthermore, our results suggest that this small remnant
population of pampas deer may be genetically differentiated from other pampas deer
populations and should be carefully managed as a separate conservation genetic unit.
(Poster presentation.)
R 97
85
DNA microsatellite analysis for parentage control of red deer in
Czech Republic.
M. Ernst
Department of Forest Protection and Game Management, Mendel University of
Agriculture and Forestry, Brno, Czech Republic
This preliminary study describe the utilization of DNA microsatellite panel for
parentage verification of red deer in Czech Republic. The efficient high-throughput
microsatellite genotyping protocol was designed by optimising microsatellite markers
already isolated for red deer ( Cervus elaphus), wapiti (Cervus elaphus canadensis),
reindeer (Rangifer tarandus f. domestica), sheep (Ovis ammon f. aries) and cattle (Bos
primigenius f. taurus). The panel is based on our assembled and present for
genotyping used set of 10 microsatellites. Randomly selected, unrelated animals of
red deer were genotyped. Genotyping was done on ABI 310 Genetic Analyser. The
results from testing of larger set of tested animals will be available in time of poster
presentation.
(Poster presentation.)
98 R
Management of
endangered deer
100 R
86
Status, ecology and conservation of barasingha (Cervus duvauceli
duvauceli) in Terai grasslands of Northern India.
J. A. Khan and A. Kaleem
Conservation Monitoring Centre, Department of Wildlife Sciences, Aligarh Muslim
University, Aligarh202 002, India
The barasingha (Cervus duvuaceli) is an endangered deer species in India. Three
subspecies of barasingha are known to occur in Indian limits. The subspecies Cervus
duvuaceli duvuaceli is found in Terai grasslands in northern India. A project was
initiated in 2003 to investigate the current population status, ecology and conservation
problems of C. d. duvuaceli. An examination of literature on past distribution of C.d.
duvuaceli coupled with extensive field surveys of extant patches of Terai grassland
showed a drastic decline in population size, distribution and abundance of this
subspecies. The C. d. duvuaceli has become locally extinct from many areas and it
currently survives at 5-6 locations where the populations are isolated and face major
conservation problems. The largest population of C. d. duvuaceli survives in Dudhwa
Tiger Reserve-Kishanpur Wildlife Sanctuary Conservation Unit (DKCU) which
approximately covers 829 km
2
. Intensive studies on abundance, habitat use, food
habits and social organization have been carried out in DKCU using a combination of
direct and indirect methods since March 2005. A total of 125 barasingha groups have
been recorded so far. The largest group comprising of 293 individuals was recorded
from Jadi Tal area in DKCU. The mean group size was found to be 35.3ยฑ5.2
(meanยฑS.E). 82% of groups had 1-20 individuals. The population in DKCU comprised
of 12.3% adult males, 14.8% sub adult and yearling males, 35.3% adult females, 9.5%
sub adult and yearling females, 16.9% fawns and 10.9% individuals could not be
classified. The mean male to female and fawn to female ratio was 50 males:100
females and 47.2 fawns:100 females. The barasingha preferred areas which had
significantly higher grass richness (t=2.4, P<0.01) and grass density (t=4.2, P<0.001).
It also utilized short grasslands more than the tall grasslands (t=16.9, P<0.001). A total
of 13 grass species were utilized by barasingha in DKCU. An analysis of various
major activities revealed that animals rested during the day time in summer. The major
threat to barasingha conservation throughout northern India is the destruction of its
habitat and illegal poaching whereas in DKCU, the illegal poaching, which takes place
when barasingha groups move out of the DKCU boundaries into agricultural fields,
emerged as major cause of its decline.
(Oral presentation.)
R 101
87
Swamp deer in Uttaranchal state, India.
S. P. Sinha
1
, S. Chandola
2
, and B. C. Sinha
3
1
Rhino Programme, C/o Wildlife Institute of India, Chandrabani Dehra Dun -248001
Uttaranchal, India
2
Forest & Wildlife Uttaranchal Forest Department, Base camp Office at Chandrabani
Road Dehra Dun- 248001Uttaranchal, India
3
Wildlife Institute of India, Chandrabani Dehradun-248001 Uttaranchal, India
Swamp deer (Cervus duvauceli duvauceli) is listed as an endangered by IUCNs Cervid
Specialist Group. The species has dwindled in number from historical levels in last
century to populations that are fragmented. One of the main reasons is the degradation
of the habitat and loss of connectivity. Dudhwa Tiger Reserve is the strong hold of
Swamp deer with population of 1250+ (2004), which also include the population in
Kishanpur Sanctuary. One small population of swamp deer is also reported in
Hastinapur in Uttar Pradesh which is under threat due to poaching, degradation of loss
of habitat due to grazing pressure and removal of grasses. All the populations are
below historical numbers and presumed carrying capacity.
Uttaranchal State which was created in 1999 is also facing problems in managing the
grassland areas along the flood plains of Ganges. Due to number of anthropogenic
pressures and developmental activities in the past and in recent years most of the
grassland areas are vanishing and in some places do not exist any more.
On 1
st
February 2005 Jhilmil Jheel was visited by me and Mr. Chandola, Adl PCCF
Wildlife, Uttaranchal Forest Department to find out the possibilities of reintroducing
rhinos in this area. While surveying the entire area with on foot and on the elephant
back we found traces of hoof marks of swamp deer. Also heard a call of a stag from
inside the tall grasses. Finally from a newly built watch tower saw 34 Swamp deer
inside an open patch of grassland along a water channel. Nine fully grown stag along
with doe and first year fawn were seen. Swamp deer antlers were also collected from
the area as evidence and for record. Jhilmil Jheel is a saucer shaped wetland situated
on the right bank of River Ganges in Chidiyapur forest range in Haridwar district in
Uttaranchal. Around Jhilmil Jheel, nine huts of Gujjars (a pastoralist community) are
situated along the forest belt and a village on the southern side. People are settled here
since 50โs. Most of them are from Punjab, Himachal Pradesh and Garhwal. In the past
there were no authentic data or information on the sighting of swamp deer in this area.
The area is rich in faunal and floral diversity considering the presence of all the five
species of deer, elephant, nilgai and tiger seen here with large number of resident and
winter migratory birds. Due to grazing pressure and presence of Gujjar community
organic matter flows directly into the Jhilmil Jheel. It has been observed that nearly
half of the Jhilmil Tall is infested by Typha and other unpalatable species. Fortunately
Pharagmities karka is flowering in the half area of the Jheel. Fortunately the local
villagers around Jhilmil Jheel are vegetarian and do not take any hard drinks because
they are followers of Sanatan dharma. After declaration of this area as Swamp deer
conservation reserve it is going to be a strong hold of swamp deer in Uttaranchal. This
102 R
area is now one of the Indiaโs first Conservation Reserve declared by Hon President
of India on 14 August, 2005. The presentation and paper is based on how this remnant
Swamp deer population is existing in this area with conservation altitude of the local
villagers and is discussed.
(Oral presentation.)
88
Distribution and abundance of wild ungulates in Royal Suklaphanta
Wildlife Reserve, Nepal.
S. Pokhrel and T. B. Thapa
Central Department of Zoology, Tribhuvan University P. O. Box: 11191 Kathmandu,
Nepal
''Distribution and Abundance of Wild Ungulates in Royal Suklaphanta Wildlife
Reserve, Nepal'' was aimed to to determine distribution, abundance and habitat
preferences of wild ungulates. The pellet groups counting along line-transect was
carried out in the western part of RSWR. A total of 7,342 pellet groups were recorded
from 2500 plots of 25 different samples. Spotted deer, hog deer, swamp deer, barking
deer, wild boar and blue bull were recorded as main ungulate species occupying the
western part of RSWR. Spotted deer was more abundantly distributed (2.28ยฑ2.23)
among all ungulate species where as blue bull was least abundant (0.002ยฑ0.05).
Ungulates were highly abundant (3.37ยฑ2.58) in grassland habitat. In four different
types of habitat, spotted deer was highly abundant (2.67ยฑ2.08) in Sal forest, hog deer
in grassland (0.53ยฑ0.74), swamp deer in grassland (1.03ยฑ1.52), barking deer in Sal
forest (0.02ยฑ0.14), blue bull in Sal forest (0.002ยฑ0.06) and wild boar in grassland
(0.13ยฑ0.034). The distribution pattern of wild ungulates was clumped type with
significant difference (P
2
= 969.28, P= 0.05 and df =24). Habitat preference was found
high in Sal forest for spotted deer (29.20%), barking deer (44.16%) and blue bull
(64.51%), and grassland for hog deer (74.81%), swamp deer (92.18%), and wild boar
(55.52%). Present study found relatively high distribution of ungulate species in core
area suggests to ungulate monitoring in extension areas.
(Oral presentation.)
R 103
89
Swamp deer (Cervus duvaceli) habitat evaluation using remote
sensing and GIS in Suklaphanta Wildlife Reserve, Nepal.
T. B. Thapa
Central Department of Zoology, Tribhuvan University P. O. Box: 11191 Kathmandu,
Nepal
Application of the remote sensing technology for wildlife habitat evaluation and
management is relatively new but, studies using remotely sensed data on physical
attributes of the habitat as well as analysis of spatial data through geospatial modelling
have been found the technology to be accurate, cost and time-effective.
I evaluated satellite derived landscape parameters to predict suitable habitat for
endangered Swamp deer (Cervus duvaceli) using GIS modelling of known wildlife
habitat relationship in the Suklaphanta Wildlife Reserve (SWR) of far western lowland
Terai, Nepal. Spatial and temporal dimension of land use/land cover was determined
by analysis of multi-temporal satellite imagery using supervised classification
techniques available in ERDAS IMAGINE 8.6 software. The pellet groups counting
along line-transect method was carried out to determine relative abundance, habitat
use and preference of the Swamp deer in the SWR. Arc GIS has been used to integrate
and analyse spatial data, in order to determine suitable habitat for Swamp deer. Six
major habitat/land-cover classes including dry sand, water body, grass land, riverain
forest, Sal forest and mixed hardwood forest were identified and delineated by
analysing Landsat ETM digital data of 2002. Habitat suitability map was prepared for
Swamp deer on the basis of linking of species with specific land cover types, water
resources and sensitivity to prevailing human activities. The swamp deer preferred
habitat mosaics consisting of grasslands, water body and small patches of forest
because such mosaic provides food, water and cover for the species. Land cover types
and their dynamics, water availability; topography, altitude, prevailing major human
activities and their impact on habitat are important landscape and anthropogenic
variables predicting suitable habitat for Swamp deer. SWR provides an excellent
habitat for a viable breeding population of Swamp deer, but the future conservation
efforts should be directed to grassland habitat management and reduction of human
intervention in the reserve.
(Oral presentation.)
104 R
90
Population Ecology of Hangul (Cervus elaphus hanglu) in Dachigam
National Park, Kashmir, India.
A. Khursheed, S. Sathyakumar, and Q. Qureshi
Wildlife Institute of India, Dehradun, India
The Hangul or Kashmir Stag (Cervus elaphus hanglu), a highly threatened species
listed in the IUCN's Red data list, is one of the four eastern most species of Red deer
(Cervus elaphus). However, unlike Red deer, which has a wide global distribution,
Hangul distribution is limited to Kashmir Himalayas in India. Today a genetically and
demographically viable population of Hangul is confined to only Dachigam National
Park (340 (34
0
05โ and 34
0
10โ to 74
0
53โ and 75
0
09โ) in the North Western
Himalayas. The present population of Hangul is estimated between 209 and 243
individuals. In this paper we made an attempt to present current information on
Hangul distribution, status and abundance along with Population Viability Analysis
(PVA). Data collection was carried out using seven line transects of varying length
(each 1- 2 km long), monitoring of trails in five fixed blocks (of 41.20 Km
2
). During
the study period (February 2001 to December 2004) a total of 693 monitoring of trails
and transects were carried out and total time and distance effort involved were 1839
hours and 5668 km respectively. A total of 326 Hangul sightings were recorded during
the trail and transect monitoring. The maximum Hangul sightings (101) were recorded
in winter and minimum (55) in summer. The overall Hangul encounter rates was 2.02
Hangul/hour effort and 0.67 Hangul/km walk and it varied significantly between
different seasons (F = 42.22; P = 0.001) and between the study blocks (F = 173.71; P
= 0.001). The overall (weighted for block areas) Hangul density recorded was 5.60
Hangul/Km
2
(S.E. = 1.13) and it varied between different seasons. There was a
significant difference mean group size of Hangul during different seasons and it
ranged between 1.10 ยฑ 0.33 in summer to 5.36 ยฑ 1.28 in spring. A typical Hangul
group size was of 14 individuals and it showed seasonal variations from 5 individuals
in summer to 17 individuals in spring. The overall Hangul sex ration was 23.23
males/100 females (S.E. = 2.60) and Hangul fawn to female ratio was 29.95
fawns/100 female (S.E. = 1.90). During four years study period the Hangul population
showed an increase at an estimated rate of 9.9% (r = 0.099). It is predicted that the
Hangul population in Dachigam National Park would show a gradual increase at an
estimated rate of 7.9% (r = 0.079 S.D. (r) = 0.129) during the next 20 years and would
reach to a maximum of 292 ยฑ 1.51 (S.E.) and after that it would remain static.
Therefore it is predicted that the Hangul population in Dachigam NP apparently is not
under any threat of extinction at least for the next 100 years. However, under the
influence of demographic stochasticity (Stoch. r.), some fluctuations in the population
growth rate are predicted in Dachigam National Park.
(Oral presentation.)
R 105
91
Microsatellite variation of Hainan Eld-s deer (Cervus eldi hainanus)
in China: Implications for conservation.
Q. Zhang, Y.-L. Song, D.-X. Zhang, and Z. Zeng
College of Wildlife Resources, Northeast Forestry University, Harbin 150040, China
Hainan Eldโs deer (Cervus eldi hainanus) experienced rapid population declined in the
end of 1960's and early 1970's. Recovery programs include in-site and of-site
conservation. in-site conservation was carried out since 1976. Effort in off-site
conservation was launched in China since 1986. The current population size is over
1300 and the demographic data may be able to sustain the population development.
However, the effect of in-site and off-site conservation on the current population
genetic structure remains to be investigated. We used 10 microsatellite DNA loci to
assess genetic differentiation and diversity for 213 Hainan Eld's deer from 6 off-site
populations. The mean number of alleles per locus was 4.1. There was significant
genetic differentiation among populations and the majority of genetic variation
(>94.65%) within populations. There was no correlation between genetic distance and
geographic distance (r = -0.1998, p < 0.8013) and there was no correlation between
genetic distance and the introduced interval time (r = -0.1093, p < 0.6010). The result
showed that population genetic variation was not completely depleted as a result of
either genetic bottlenecks or founder events due to the deer population had a certain
extent genetic variation and past management and conservation actions are successful
for populations maintaining fine-scale genetic structuring. Based on significant genetic
differentiation among Hainan Eld's deer populations, where possible, it may be
advisable to translocate individuals between isolated populations to maintain levels
of genetic variation in remaining Hainnan Eld's deer populations.
(Oral presentation.)
92
Social structure of the reintroduced Persian fallow deer (Dama
mesopotamica) population: integrating three observation methods.
A. Perelberg, S. Bar-David, U. Roll, A. Dolev, and D. Saltz
Mammals Research Center, the Society for the Protection of Nature in Israel (PA,
DA), Israel
In order to increase the probability of success of reintroductions, it is essential to
estimate the social-structure of the reintroduced species. Collection of this information
is especially difficult in hard-to-observe species (e.g., hidden in the thicket).
Additionally, previous knowledge about many reintroduced species is unavailable due
to their rarity in their original environment. We studied the social-structure of the
106 R
Persian fallow deer population, one of the rarest deer in the world, now being
gradually reintroduced to the Western Galilee, Israel, since 1996. The deer were bred
in captivity under artificial conditions for many generations, and we feared that they
will not be able to adjust their social-structure as expected from a wild population. We
used three observation methods: 1) radio-telemetry collars mounted mostly on females
(due to the technical difficulties of tagging males); 2) direct-observations, conducted
from pre-defined observation points and occasional sightings along roads; and 3)
portable IR video-camera, installed in deer activity areas. We found that in the winter,
groups were larger and associations were stronger than in other seasons. Groups were
also larger at night. Adult males were in smaller groups than young males and
females. Additionally, negative correlations were found between group-size to both
time form release and distance from release site. Group size stabilized in ca. 6 months
after release on 1.670.9 (meanSD), with a tendency of adult males to be solitary.
Overall, the deer apparently do not keep fixed social-structure over time, even when
home-ranges highly overlap. Integrating the three methods provided us with a more
accurate picture of the deer social-structure: telemetry provided continuous recordings
of home-ranges overlap and assessment of association levels, whereas the other
methods were more sporadic. However, because most reintroduced males and wild-
born fawns were not collared, a complete assessment of the social-structure was
impossible without the implementation of these additional methods. Video-recordings
added important information about the social-structure at night and in the thicket, that
could not be otherwise obtained. Together with already reported parameters, it seems
that the deer established a viable wild population, and that the chances for a successful
reintroduction are high.
(Oral presentation.)
93
Ecology and conservation of the huemul in southern Chile.
R. Gill
1
, C. Saucedo
2
, and D. Aldridge
3
1
Ecology Division, Forest Research, Alice Holt Lodge,Wrecclesham, Surrey GU10
4LH, UK
2
Departamento Patrimonio Silvestre Corporaciรณn Nacional Forestal (CONAF), /
Center for Andean Wildlife Research (CAWR) VIII Regiรณn, Chillรกn Chile.
3
Departamento Patrimonio Silvestre Corporaciรณn Nacional Forestal, XI Region, Avda
Ogana 1060, Coyhaique, Chile.
The huemul Hippocamelus bisulcus is an endangered deer occupying temperate
woodland habitats in the Andes of southern Chile and Argentina. The distribution and
numbers have declined substantially since settlement by Europeans, due mainly to a
combination of hunting and habitat loss. However, current information on ranging
behaviour and population ecology in general is very limited. We selected three sites
selected in Aysรฉn, Chilean Patagonia, to study the movements, survival and habitat
R 107
associations of huemul using radio telemetry. Although substantial seasonal
migrations are commonly reported amongst other deer species in mountainous regions,
we found that huemul undertook either no or only modest movement between summer
and winter. Long distance movements (>5km) were very infrequent. Habitats selected
by huemul included lenga Nothfagus pumilio forest, rocky places and open shrubs;
those avoided included grassland and steppe. Amongst the sample of marked animals
and their offspring, we found that recruitment rates were barely sufficient to balance
mortality. The principle sources of mortality included poaching and predation by
puma, dogs and foxes (on fawns). Although huemul are protected in a number of
reserves in both Chile and Argentina, populations are thinly distributed in suitable
habitat along valley sides and there is little scope for increasing the number of
protected areas. Conservation efforts are currently focussed on raising awareness of
the plight of the huemul amongst farmers and the general public, and encouraging
sympathetic land use practices, such as ecotourism.
(Oral presentation.)
94
Status, genetic structure and Conservation suggestion of Chinese
water deer.
M. Chen and E. Zhang
East China Normal University, Life Science School, Shanghai, 200062, China
Chinese water deer (Hydropotes inermis inermis) is an endangered species, ranked as
LR/nt in IUCN red data book, which is native to China. The distribution of the water
deer in China used to extend to Liaodong PeninsulaยฃยฌNorth China plain and the
shores of the lower reaches of Yangtze River and adjacent lake areas, with a region
from the latitude 28
o
to 42
o
N and the east limit, the longitude 110
o
E. However, due
to habitat loss and hunting, the population of the water deer has declined rapidly and
its distribution is now limited and fragmented only to the coastal areas of Jiangsu
Province, Zhoushan Archipelago of Zhejiang Province, and Poyang Lake areas in
Jiangxi Province. In order to decide conservation strategy, we analysis the reason of
the deerโs population shrink sharply and do test to understand the genetic structure of
the distributions. In view of climate variations in the past 10,000 years and human
population growth, We consider that the deerโs population changes in North China
was mostly due to climate change, agriculture development, human population
increase and vegetation degradation. The populations of Chinese water deer shrunk
sharply in China recently, especially the lower reaches of Yangtze River and its
adjacent lake areas and south of Yangtze River. The reasons were no others but
habitats lost and poaching.
The cytb, D-loop and 12SrRNA of mtDNA were sequenced from the samples
collected noninvasively from Zhejiang, Jiangsu and Jiangxi province. The results
show that Chinese water deer have high genetic diversity and the population in
108 R
Jiangsu is the highest. The population from Jiangxi should be ESU and worthy of
conservation consideration. There are no clear diverging between the populations from
Zhejiang and Jiangsu province. Another conservation suggestion is reintroducing this
animal to the area where Chinese water deer existed before to enlarge and restore the
population area where Chinese water deer existed before to enlarge and restore the
populations.
(Oral presentation.)
95
Spatial pattern characteristics of wapiti habitat fragmentation
factors based on spatial autocorrelation and semi-variance analysis
in Northeastern China.
M. Zhang, G. Jiang, and J. Ma
College of Wildlife Resources, Northeast Forestry University, Harbin 150040, China
Both spatial autocorrelation analysis and semi-variance analysis by Geo-Statistics
were used to study the spatial pattern characteristics of wapiti Cervus elaphus
xanthopygus Milne-Edwards' habitat fragmentation factors during three winters from
2003 to 2005, which revealed the relationship between patch scale structure and wapiti
population distribution. Our data suggested that: 1) Under fourteen distance classes
(2303220 m), nine habitat fragmentation factors had spatial autocorrelation
significantly (P < 0.05) and Moran-s I reached at 73%; Six habitat factors appeared
different spatial correlation characteristics significantly under different distance
classes, which indicated that there were different microhabitat scale structure
characteristics of these habitat factors. Three habitat factors (Settlements, abandoned
logging roads and farmlands) appeared correlated significantly under all fourteen
distance classes, which indicated not only higher consistency in influencing on the
spatial pattern of habitat fragmentation, but also their important role in the process of
habitat fragmentation of wapiti during winter; 2) Semi-variance analysis and fractals
analysis showed that nest structure characteristics of the changes of semi-variance of
population distribution responding to the changes of spatial distance classes, which
revealed that there was a multiple scale variations in spatial pattern of wapiti
distribution; however, fractal number-D of wapiti distribution approached 2, which
indicated the variation of spatial pattern of wapiti distribution mainly occurred not
only at smaller scale but larger scale for larger standard deviation, which showed that
differences of track numbers of wapiti occurred in sample sites were very large, and
small numbers came after a period time of larger numbers of wapiti tracks. The above
data showed that multiple scale of wapiti distribution, class patches of spatial
distribution and vulnerability of wapiti microhabitats, indicating that spatial pattern
of wapiti distribution would change greatly once microhabitat changed. In addition,
it was of importance for validity, independence and representative of applying survey
R 109
scale during wapiti habitat research through revealing spatial autocorrelation and
distribution patterns of habitat fragmentation factors under different spatial scales in
the future.
(Oral presentation.)
96
Assisted reproductive technologies for endangered deer species.
Y. Locatelli, J.-C. Vallet, X. Legendre, and P. Mermillod
Rรฉserve animaliร re de la Haute Touche, Musรฉum National d'Histoire Naturelle,
36290 Obterre, France
INRA Physiologie de la reproduction et des comportements, Equipe follicule Ovocyte
et Dรฉveloppement, 37380 Nouzilly, France
Within two hundred cervid sub-species over the world, almost 40 subspecies are
considered to be threatened with extinction by the IUCN (International Union for
Conservation of Nature and Natural Resources). In most of the cases, preservation of
this biodiversity relies on conservatory programs performed in and ex situ. In parallel
with ex situ conservatory programs based on classical farming in zoo, assisted
reproductive technologies may represent an efficient way to produce and disseminate
offspring and to prevent genetic loss from rare remaining individuals. In this aim,
association of techniques such as recovery of semen/oocyte and in vitro production of
embryos appears especially suitable for deer species. Achievability of in vitro embryo
production methods was investigated for Cervus nippon and elaphus after processing
of oocytes recovered respectively by aspiration of follicles from live donors using
laparoscopic ovum pick-up (LOPU) after gonadotrophin stimulation (0,5 IU oFSH)
during the breeding and non breeding season or from abattoir-derived ovaries during
the breeding season. LOPU collection from live sika deer hinds were performed with
minimal stress to the donor and were repeated without complication. An average of
8 to 10 cumulus-oocyte complexes (COC) were collected per hind per LOPU session
during the breeding season vs. 4 to 5 COC during the non breeding season. After in
vitro maturation and fertilization, about 60 to 80 % of sika or red deer cumulus-oocyte
complexes were fertilized. Developmental competence was assessed for both red deer
and sika deer embryos in two different culture systems. Embryos were cultured in
synthetic oviduct fluid (SOF) medium in the presence or absence of ovine oviduct
epithelial cells (oOEC). In contrast with classical SOF culture, high proportion of
embryos reached the blastocyst stage in presence of oOEC (20-40 % with oOEC vs.
0-5 % in culture medium alone). Viability of frozen red deer embryo produced in vitro
on oOEC was assessed by transfer of ten blastocysts to five synchronised recipients.
Three male calves (one from vitrification and two from slow freezing) were born
unassisted after 237 to 238 days of gestation (Locatelli et al., 2005). These results
illustrate achievability of IVP procedures in two deer species and may offer new
possibilities for conservation of rare deer species in future.
110 R
(Oral presentation.)
97
Diet composition and habitat selection of red deer during winter in
Helan Mountains, China.
Z. S. Liu and X. M. Wang
School of Life Science, East China Normal University, 3663 Zhongshan N. Rd,
Shanghai 200062, China
Diet composition of red deer in Helan Mountains of China was determined by fecal
and rumen content analysis from November 2003 to February 2004. Habitat
characteristics were measured by locating from fresh sign or dung on line transects.
Red deer consumed 50 plant species, including 9 woody species, 14 shrubs, 5
graminoids, 2 sedges, 19 forbs and other plants unidentified. Woody species, such as
Populus davidiana and Salix spp. were utilized significantly more than in proportion
to availability. Of the shrubs, Ribes spp., Dasiphora spp., Caragana spp., Syringa
oblata, and Lespedeza spp. were selected in greater proportion than its availability.
Graminoid species used significantly more than expected included Tragus
mongolorum, Calamagrostis spp., and Echinops spp.. Forb species, including Urtica
spp. and Thymus spp. were used significantly more than available. Deer preferred
montane grassland and montane conifer forests, and used montane savanna in
proportion to its availability. Red deer avoided subalpine shrublands and meadows.
Red deer preferred mixture habitats, and avoided habitats dominated by Juniperus
rigida and Pinus tabulaeformis. Deer showed a strong preference for slope exposed to
the sun, and avoided shady slope. Red deer preferred lower slope, and avoided upper
and middle slope. Compared with random plots, usage sites were characterized by
more shrubs, higher food abundance, lower slope, shallower snow cover and approach
to water. Most sites were located farther away from bare rock and human disturbance.
Results of principal component analysis showed that the first 6 principal components
explained 84.89% of the total variance among all habitat factors. PCA indicated that
vegetation type, tree density, shrub density, slope degree, slope direction, snow depth
and distance to water were important factors in habitat selection.
(Poster presentation.)
R 111
98
Conservation status quo and study progress of Siberian musk deer
(Moschus moschiferus) in China.
J. Wu and Y. Zhang
College of Wildlife Resoures, Northeast Forest University, Harbin 150040, China
In China, Musk deer mainly distribute in Heilongjiang, Jilin, Mongolia provinces etc.
It is national degree T conservation animal, listed in CITES appendix U. Recently, no
more than 4000 musk deer are reserved in China, and the number is still reducing that
caused by illegal capture for economic benefits and lost of suitable habitats. Mountain
close for plantation, natural forest protection project and plenty of wildlife nature
reserve establishment protect the habitat of musk deer effectively. In addition, a series
of legislation were established to protect wildlife and reinforce the conservation of
musk deer. The habitat selection and food composition of musk deer was investigated
in Huzhong and Tonghe of Heilongjiang province since 2004 following some results:
musk deer prefer to the coniferous broadleaved forests, frequently inhabit in the plots
that full of cliff bid stone. The habitats are characterized by high altitude, small
canopy, abundant food and steep slope. The diet consist of 84 plant species of 56
families, including 33 wood plants (39.28%), 23 herbages (27.38%), 16 mosses
(19.05%), 6 ferns (7.14%), 6 lichens (7.14%). The musk deer mainly graze the tip of
the plants, such as twig, wand and tender. The mostly food is the twig of wood plants
for the whole year especially the low shrub. The wood plants consist of Rhododendron
dauricum, Acer mono, Deutzia glabata, Populus ussuriensis and Tilia amurenssis etc.
and the herbage consist of Carex callitrichos and Caulophyllum robustum etc.
(Poster presentation.)
99
Agonistic and non-agonistic behaviour interactions in Indian
blackbuck (Antelope cervicapra L.) during dominance hierarchy
formation.
T. Rajagopal and G. Archunan
Research Scholar, Bharathidasan University, Department of Animal Science,
Tiruchirappalli-620 024, India
The limited availability of necessary resources, such as food, shelter, space and mating
opportunities (i.e. selection of female for mate), provokes an agonistic and non-
agonistic interaction that lead to the establishment of dominant hierarchies. These
interactions may leads to fighting, chasing and the formation of dominance hierarchy
112 R
that determines the order of access to both present and future resources. The present
investigation was carried out at Arignar Anna Zoological Park, Vandalure, Chennai,
to assess the frequency and duration of agonistic (i.e. fighting, chasing and scent
marking) and non-agonistic behaviours (i.e. resting and feeding), before, during and
after the formation of dominance hierarchy (i.e. determine the territorial ownership)
in blackbuck herd. We selected two types of blackbuck groups i.e. dominant and
subordinate males. Behaviour was recorded by focal sampling method at intervals
through the study period. During field observation, before hierarchy formation the
dominant buck spent more time in offensive display, while subordinate buck spent
more time in defensive locomotion, although offensive display was strongly affected
by subordinate buck for formation of higher social status. Moreover, during hierarchy
formation the dominant buck showed higher frequency of scent marking, reproductive
and feeding behaviours as compared to subordinate buck followed by before and after
hierarchy formation. After the hierarchy was formed, if dominant buck becomes
subordinate, then it showed very little aggressive behaviour and spent much more time
on resting and submissive behaviour. The occurrence of agonistic and non-agonistic
behaviours in blackbuck herd clearly indicates that before and during dominance
hierarchy formation, the fighting and chasing behaviours are inflicted injuries in
subordinates. The results suggest that dominant male scentmarking odours
(pheromonal cues) may rely on strategies to suppress the aggression, mating, scent-
marking, scent production, territorial patrolling activities in subordinate bucks,
through which the dominant male exhibits its hierarchy and success in reproduction.
(Poster presentation.)
Reproduction
114 R
100
Gossypol-based contraception in male deer (Cervus elaphus).
Z. Giลผejewski
1
, B. Szafranska
2
, Z. Steplewski
3
, G. Panasiewicz
2
, and H. Koprowski
3
1
Polish Academy of Sciences, Institute of Animal Reproduction and Food Research,
PL-10-747 Olsztyn, Poland
2
University of Warmia and Mazury, Department of Animal Physiology, PL-10-719
Olsztyn-Kortowo, Poland
3
Biotechnology Foundation Laboratories, Thomas Jefferson University, Philadelphia,
PA 19107, USA
Semen samples from 4 adult red deer (Cervus elaphus) fed cottonseed (350g for 109
days) as a source of gossypol (15mg/kg body weight/day) were analysed quantitatively
and qualitatively by classical methods used in domestic animals. The data from 182
ejaculates were compared with those obtained from 571 ejaculates of stags (n = 5)
during three previous natural reproductive seasons. Spermatozoa of mature male deer
fed cottonseed exhibited an increase in morphological abnormalities (strongly folded
tails) and a decrease in motility, and concentration, leading to decreased semen quality
but no detectable side effects in the animals. Follow-up during the next reproductive
season when gossypol ingestion was discontinued revealed a return to baseline values
for average sperm concentration and motility. These findings point to the influence
and safety of gossypol fed to deer in the form of cottonseed as an efficient male
contraceptive.
(Oral presentation.)
101
The hoarse vocalization and the inflatable laryngeal air sac of
reindeer (Rangifer tarandus).
R. Frey, A. Gebler, G. Fritsch, K. Nygrรฉn, and G. E. Weissengruber
Institute for Zoo and Wildlife Research (IZW), Alfred-Kowalke-Strasse 17,
10315 Berlin, Germany
The vocal tract of reindeer has evolved an inflatable ventrorostral laryngeal air sac.
This air sac is not present at birth but emerges during ontogenetic development. It
protrudes from the laryngeal vestibulum via a short pneumatic duct. In the female the
growth of the air sac stops at an age of 2-3 years whereas in males it continues to grow
up to an age of about 6 years leading to a pronounced sexual dimorphism of the air
sac. In both adult females and males the ventral air sac walls touch the integument. In
the adult male the air sac is laterally covered by the mandibular portion of the
sternocephalic muscle and the skin. Both sexes of reindeer have a double stylohyoid
R 115
muscle and a thyroepiglottic muscle. These muscles may assist in inflation of the air
sac. Head and neck specimens were subjected to macroscopic anatomical dissection,
computer tomographic analysis, and skeletonization. Acoustic recordings were made
during a fall round-up of semi-domestic reindeer in Finland and in a small zoo herd.
Male reindeer adopt a specific posture when emitting their serial hoarse rutting calls.
Head and neck are kept low and the throat region is extended. In the ventral neck
region, roughly corresponding to the position of the large air sac, there is a mane of
longer hairs. Neck swelling and mane spreading during vocalization may act as an
optical signal to other males and females. The air sac, as a side branch of the vocal
tract, can be considered as an additional acoustic filter. Individual acoustic recognition
may have been the primary function in the evolution of a size-variable air sac and this
function is retained in mother-young communication. In males sexual selection seems
to have favoured a considerable size increase of the air sac and vocalization became
restricted to the rutting period serving the attraction of females. We propose two
possibilities for the acoustic function of the air sac in vocalization that do not exclude
each other.
(Oral presentation.)
102
Patterns of long-term reproductive success in male and female
white-tailed deer.
R. W. DeYoung
1
, K. L. Gee
2
, S. Demarais
3
, R. L. Honeycutt
4
, and R. A. Gonzales
2
1
Caesar Kleberg Wildlife Research Institute, Texas A&M University-Kingsville,
Kingsville, TX 78363 USA
2
Samuel Roberts Noble Foundation, Ardmore, OK 73401 USA
3
Departent of Wildlife & Fisheries, Mississippi State University, Mississippi State, MS
39762 USA
4
Department of Wildlife & Fisheries Sciences, Texas A&M University, College
Station, TX 77843 USA
Female white-tailed deer (Odocoileus virginianus) occur in small groups dispersed
throughout brushy habitats. Male mating tactics such as harem holding, resource
defense, etc., are not profitable in these environments, so males use a wandering
strategy to locate individual estrous females. Although ecological and behavioral
factors are known to affect the reproductive success of males, there are few long-term
studies of reproductive success in ungulate species. We used genetic parentage
assignments over 11 annual fawn cohorts to investigate the variance in long-term
reproductive success (LRS) of male and female white-tailed deer on a 1,200-ha
management area in Oklahoma, USA. Estimates of LRS during the study period
indicate that for successful breeders, male LRS was higher and more variable than that
of females (male mean = 2.5, var = 6.2, maximum = 12; female mean = 1.7, var = 1.1,
maximum = 7). Reproductive success was strongly age-dependent for both sexes, with
116 R
86% and 67% of offspring assigned to females and males ร3.5 years old, respectively.
Breeding lifespan (L) was an important determinant of female LRS, but variance
components of L and annual fecundity (F) covaried negatively, indicating that the
advantages of living longer could be offset through variable fecundity among years.
Circumstantial evidence indicates that fawn survival also is an important factor in
female LRS, since few females successfully recruited twin fawns within years or
single fawns in successive years. These observations may reflect the importance of
adequate nutritional reserves or maternal experience in fawn recruitment. Variance
components for male L and F were similar, suggesting that males could achieve high
LRS either by living longer or through increased F. A positive covariance between
male L and F is consistent with greater F for older males. This study, the first to
document LRS in white-tailed deer, emphasizes the differential effects of behavioral,
ecological and life history variables on fitness between the sexes. The reproductive
success of individual males is limited and breeding is distributed among a greater
number of males in this mating system. As a result, the difference in LRS between the
sexes in white-tailed deer is much less than in other ungulate species, which are highly
polygynous.
(Oral presentation.)
103
Observations on the reproductive behaviour of sambar deer (Cervus
unicolor unicolor) in a bush enclosure in Victoria, Australia.
W. M. Harrison, I. A. Moore, M. Draisma, and G. I. Moore
Australian Deer Research Foundation Ltd., Australia
The reproductive behaviour of sambar deer (Cervus unicolor unicolor) was monitored
in a 13ha forested enclosure in Victoria over a twenty-year period during which time
sixty-one calves were born.
Antler casting was relatively synchronised in stags, but variable in timing from year
to year and was recorded from November through February, peaking during January.
Antlers were normally hard and cleaned of velvet in May-June. Signposting by stags
comprised antler rubbing, wallowing, scraping and preaching and commenced from
the time antlers were cleaned of velvet through until just prior to casting. These
activities were recorded from May through December, peaking in September. The
activity most frequently recorded was wallowing, with stags as young as five-months-
of-age taking part. Wallowing, particularly in the dominant stag, was clearly sexually
related. Changes in antler status (antler casting) influenced a temporary change in
dominance rank. A dominance ranking in both males and females was evident. An
oestrus cycle of about twenty days appears likely and the data was suggestive of a
gestation period of about 260 days. The mean age at which seven hinds first conceived
was 16.3 months (Range 14.3-20.1), and parturition of first calves occurred at a mean
age of 24.9 months (Range 22.8-28.6). One eighteen-year-old hind born in the
enclosure reared fourteen calves, another, a yearling when introduced, was known to
R 117
have borne twelve calves when, at the age of fifteen years, she escaped from the
enclosure. The length and timing of breeding and calving seasons of sambar in the
wild has been the subject of conjecture in the literature. Based on a 260-day gestation
period, we found conceptions to have occurred in all months excepting February,
March, and December, with peak breeding activity taking place from June through
November. One conception occurred during January, in a year in which stags did not
cast antlers until February. Two hinds conceived when stags were in late stages of
antler growth. Births peaked from March through July with none occurring from
October to December, nor during February.
(Oral presentation.)
104
Sexual choice in lekking fallow deer (Dama dama): variable female
strategies.
S. Imperio, S. Focardi, F. Ronchi, and A. M. De Marinis
Istituto Nazionale per la Fauna Selvatica, Universitร di Firenze, Italy
Here we report one of the first analysis of female mating strategies in lekking fallow
deer. The present work try to answer two basic questions. Do females use different
strategies of mate selection at lek? If so, do such strategies depend on costs and/or
experience of females?
We studied the behaviour of 10 radio-collared and 92 ear-tagged fallow deer females
during 4 rutting periods (2000 to 2003) in the Preserve of Castelporziano, Rome. Each
female was assigned to a distance class (distance between home range and lek) and
this was assumed to be a proxy for the cost of moving to the arena. Since most of
females use to visit the lek every year, their age covaries with experience. Younger
females spend more time at lek and mate significantly later than adult ones. We
interpret the observed increase in the variance in male reproductive success observed
after the peak of the rut, as determined by an increase in the unanimity of female
choice because young fallow does copy the mate choice of adults. Females living
close to the lek, begin to visit the arena early in the rutting period where they spend
much time (up to 208 hours), maybe to have the opportunity to assess the quality of
the largest number possible of bucks. On the contrary, females living farther from the
arena visit males at lek only the time strictly necessary for mating. To increase the
precision of choice, these females make use forms of indirect assessment of male
quality, such as association to other females and selecting territories which were
characterised by high male success in the precedent year, but they do not seem to copy
the choice of other females.
Females living at more than 8 km from lek usually do not to visit the arena, since costs
for moving are probably too high. Besides, we demonstrate that non-independent
strategies of male quality assessment do not results in the same benefits of direct
assessment.
118 R
(Oral presentation.)
105
Variation in fawn production in a semi arid environment: An
energetics approach.
D. G. Hewitt and E. L. Monaco
Texas A&M University-Kingsville and the Caesar Kleberg Wildlife Research Institute,
Kingsville, Texas 78363, USA
Fawn recruitment is a critical demographic parameter in management of deer
populations. In semi-arid southern Texas, USA, fawns are born in June and July when
forage production are influenced heavily by heat and precipitation. Thus, fawn
recruitment is positively correlated with spring-summer rainfall. Whether low fawn
recruitment during drought is the result of increased predation from poor hiding cover,
poor forage quality for the doe, or a combination of both, is unknown. Understanding
the relative effects of predation and nutrition on recruitment is important in designing
management programs to increase fawn recruitment. For example, some management
programs seek to increase grass cover, which provides additional hiding cover at the
expense of high quality forage. To test the idea that forage quality could limit
reproduction in a dry year, we modeled energy intake and energy expenditure of a doe
from breeding in December through fawn weaning in September. The model tracked
the does fat reserves and when her body fat dropped below 5%, we assumed she was
in too poor condition to finish raising her fawns. Diet quality and initial doe fat
reserves were varied to understand their interactions. A doe consuming a poor quality
diet, such as that expected when consuming browse, could not produce fawns, even
if the doe had large fat reserves in December. A deer consuming a high quality diet
was able to raise fawns even if she was in poor condition when she was bred. On an
annual diet consumed by deer in South Texas (high quality in late winter, moderate
through the spring, and poor in July and August), deer needed to enter the
reproductive season in good body condition to successfully produce fawns, otherwise
they mobilized all their fat reserves before their fawns were weaned. Our model
suggests that poor fawn production during dry summers is caused in part by nutritive
failure. Management projects that seek to increase fawn cover at the expense of high
quality forage plants could be detrimental.
(Oral presentation.)
R 119
106
Movements of female white-tailed deer during parturition and the
rut in a high-quality, balanced sex ratio herd in Maryland, USA.
L. I. Muller
1
, K. A. Adams
2
, M. C. Conner
3
, and J. L. Bowman
4
1
University of Tennessee, Dept. Forestry, Wildlife and Fisheries, Knoxville, Tennessee
37996, USA
2
National Wild Turkey Federation, USA
3
Dupont Agricultural Products, USA
4
University of Delaware, USA
White-tailed deer (Odocoileus virginianus) movements are affected by life history,
density, and demographics. Many white-tailed deer herds have low numbers of males
because of past harvest strategies. We examined deer movements during peak life-
history events in high-quality deer habitat with balanced sex ratios (approximately
1:1.5) and overall deer density of 25-30 deer/km
2
. This study was conducted on
Chesapeake Farms (Kent County, Maryland, USA). We attached 16 GPS-telemetry
collars to does during the spring of 2001 (n = 10) and 2002 (n = 6). Seven does were
also fitted with vaginal-implant transmitters to monitor fawning (n = 6 and 1 for 2001
and 2002, respectively). Twelve collars collected useable location data pre- to post-
parturition. Two collars collected data during autumn. Mean adaptive kernel summer
home-range size was 32.3 ha. Mean parturition date was 28 May (27 May 3 June).
Mean 4-hr distances (straight-line distances between 4-hr locations) for mature does
10 days prior to parturition, 10 days surrounding parturition, 1-10 days after
parturition and 11-20 days after parturition were 202.4 m (SD = 238.8), 131.2 m (SD
= 159.3), 104.5 m (SD = 94.7), and 154.5 m (SD = 141.9), respectively. Does
restricted movements during parturition and shortly afterwards; however, they
resumed normal movements after 10 days post-birthing. Female white-tailed deer have
been shown to exhibit 2 different behaviors during breeding depending on male
density. Females reduced movements to ensure predictable locations with high male
density. With low male density, does increased movements to actively search for
mates. The 2 collared does exhibited movements indicative of an active search
strategy even though males were not limiting. The 5.5 year-old doe in 2001 increased
her daily mean 4-hr movements from 320.0 m (SD = 260.4) on 11 November to 966.1
m (SD = 844.7) on 12 November and returned back to lower activity on 13 November
(200.3 m, SD = 222.3). These movements were across a tidal creek and outside of her
normal home range. The 3.5 year-old in 2002 exhibited a similar pattern. On 8
November, her daily mean 4-hr movements were 166.6 m (SD = 166.5). She increased
these movements on 9 November (368.0 m, SD = 408.0) and returned to lower
movement (130.5 m, SD = 183.1) on 10 November. We were able to observe these
movements with the GPS technology and they corresponded to peak rut dates based
on fawning data. Potential active search strategy by does in an area with high male
density may indicate female mate choice, provide a mechanism for inbreeding
avoidance, and enhance reproductive success.
(Oral presentation.)
120 R
107
Refrigerated storage impairs chromatin of Iberian red deer (Cervus
elaphus hispanicus) epididymal spermatozoa kept inside the
epididymis.
A. E. Dominguez-Rebolledo, M. C. Esteso, M. R. Fรฉrnandez-Santos, D. Matias, F.
Martinez-Pastor, and J. J. Garde
Instituto de Investigaciรณn en Recursos Cinegรฉticos, IREC (CSIC-UCLM-JCCM),
Albacete, Spain.
Postmortem time negatively affects seminal samples from the cauda epididymis, a
potential source of spermatozoa for germplasm banks, and little is known about its
effect on sperm DNA. We have studied the effect of chilled storage of genitalia on the
chromatin of epididymal spermatozoa, obtained from the cynegetic species Iberian red
deer (Cervus elaphus hispanicus). Testicles were collected from 29 stags harvested
during the hunting season. The testicles (5 h postmortem) were put into tightly closed
plastic bags, placed in a beaker with water and then stored at 5ยบC. At 0, 24, 96 and 192
h, sperm was extracted from the cauda epididymis by means of cuts. Chromatin was
assessed by flow cytometry after detergent-acid (pH 1.4) treatment and acridine
orange staining. We obtained the parameters SDat (sperm heterogeneity regarding
chromatin status), DFIm (% spermatozoa with moderate DNA Fragmentation Index)
and DFIh (% spermatozoa with high DNA Fragmentation Index). The effect of storage
time was analyzed by linear mixed-effects models, including male as random effect,
and time as fixed effects (covariate). The models showed and increase in all the
parameters, indicating that storage time negatively affected chromatin status (P<0.01
for SDat and mDFI, and P<0.1 for hDFI). Thus, results at 0 h were (meanยนSD): SDat:
2.1ยน1.4: DFIm: 0.4ยน0.6%; DFIh: 0.1ยน0.2%; whereas at 192 h, results were: SDat:
3.3ยน1.6: DFIm: 1.6ยน1.5%; DFIh: 0.2ยน0.3%. In conclusion, red deer epididymal
spermatozoa underwent chromatin damage during chilled storage inside the
epididymes. Nevertheless, the impairment were not very important, therefore
epididymal samples may be stored inside the epididymes for a long time if necessary,
being still usable for IVF or ICSI techniques. However, this situation should be
avoided whenever possible.
Supported by grant AGL2004-05904/GAN from Ministerio Espaรฑol de Educaciรณn y
Ciencia (MEC). F. Martรญnez-Pastor is supported by the Juan de la Cierva program
(MEC).
(Poster presentation.)
R 121
108
Immunohistochemical expression of steroidogenic enzymes in the
corpus luteum and placenta of sika deer (Cervus nippon) during
pregnancy.
Y. Matsuura
1
, D. Hayakawa
2
, Y. Yanagawa
1
, M. Sasaki
2
, H. Igota
3
, C. Yayota
1
, S.
Kondo
1
, N. Kitamura
2
, T. Tsubota
4
, and M. Suzuki
1
1
Hokkaido University, Graduate School of Veterinary Medicine, N18-W9, Kita-ku,
Sapporo, Hokkaido 060-0818, Japan
2
Obihiro University of Agriculture and Veterinary Medicine, Japan
3
Nisiokoppe Wildlife Association, Japan
4
Gifu University, Japan
Multiple (usually two) corpora lutea (CLs) are often found in the ovaries of sika deer,
although they basically deliver a single fawn. Since the corpus luteum (CL) is the
major structure that secretes progesterone, we presume that the existence of multiple
CLs affects the achievement and maintenance of pregnancy. In this study, we
investigated the expression of steriodogenic enzymes, i.e., cholesterol side-chain
cleavage cytochrome P450 (P450scc), 3รข-hydroxysteroid dehydrogenase (3รขHSD),
aromatase cytochrome P450 (P450arom) and 17รก-hydroxylase cytochrome P450
(P450c17), to compare the process of steroids production and the localization of
immunostaining cells between two CLs found in individual animals during pregnancy.
The placenta also synthesizes progesterone; however, the synthesis pattern differs
depending on the species. Then, we also applied the same analysis to placenta in sika
deer. Ovaries, placentae and fetuses were collected from pregnant female sika deer
which had been killed for research during pregnancy periods (i.e., from December to
May). Fetuses were used to estimate the gestation ages. Almost all luteal cells of both
CLs expressed P450scc, 3รขHSD, P450arom and P450c17 throughout pregnancy.
During mating season, only luteal cells located in the periphery of the CL showed
positive immunostaining in latter formed CL. However, both CLs were similar in
terms of their immunostaining pattern during pregnancy. There were small luteal cells
(thecal cells) and large luteal cells (granulosal cells), and both reacted similarly at least
until middle stage of gestation, when small luteal cells could be rarely observed.
Regarding the placenta, immunoreactions of P450scc, 3รขHSD, P450arom and
P450c17 were observed mainly in the chorionic villi at all gestational stages studied
and tended to be strong in trophoblast giant cell. These results suggest that throughout
pregnancy, both CLs have similar capabilities for synthesizing steroid hormones. And
also, the placenta as well as the CL has probably the ability to synthesize
progesterone, estrogen, and androgen throughout pregnancy, and both organs may
play an important role in the maintenance of pregnancy in sika deer.
(Poster presentation.)
122 R
109
Objective quality control of frozen-thawed red deer spermatozoa by
Computer-Assisted Semen Analysis - instrument settings.
Sz. Nagy
1
, E. Puskรกs
2
, I. Pรฉntek
3
, and Z. Zomborszky
2
1
Research Institute for Animal Breeding and Nutrition, H2053 Herceghalom,
Gesztenyรฉs str.1., Hungary
2
University of Kaposvรกr, Faculty of Animal Sciences, H7400 Kaposvรกr, Guba S. str.
40., Hungary
3
National Institute for Agricultural Quality Control, H1024 Budapest, Keleti Kรกroly
str. 24., Hungary
Cryopreservation of spermatozoa as a tool for genome resource banking of wild
animals is well known. Moreover, artificial insemination with frozen-thawed sperm
is already used in breeding of some farmed wild species like deer. Computer-Assisted
Semen Analyzers (CASA-s) are useful for the quality control of frozen-thawed sperm
as they are able to measure several parameters at cellular level and on a high number
of cells; however, they require different instrument settings for the different species.
The aim of the present study was to establish a red deer-specific CASA setting.
During the rutting season of 2004, epididymal sperm samples of five red deer (Cervus
elaphus hippelaphus) stags were collected post mortem in South-west Hungary.
Samples were processed in a commercial bovine semen extender (BioXcell, IMV,
LAigle, France), filled into 0.25 ml straws then they were frozen by holding in liquid
N
2
vapor. Final cell concentration was adjusted to 100 x 10
6
/ml. Initial motility was
assessed visually under a phase contrast microscope at 200x magnification. For the
post-thaw quality control by CASA (Medealab, MTG, Altdorf, Germany), sperm
samples were thawed by holding the straws in a 38
o
C water bath for 10 sec, diluted
to approximately 50 x 10
6
/ml with BioXcell, then measurements were taken on 12
fields per sample, using a 20x Negative High Contrast objective and a 20 ยตm deep
counting chamber and a heated microscopic stage. Species specific Average Path
Velocity (VAP, ยตm/s) threshold values of the different categories (nonmotile, slow
progressive, fast progressive spermatozoa) were calculated by k-cluster analysis
(Statistica for Windows 6.0).The VAP threshold values revealed by k-cluster analysis
were the following: Nonmotile: <10 ยตm/s; Slow progressive: >20 ยตm/s; Fast
progressive: >80 ยตm/s. With the help of this objective quality control method, our
laboratory is ready to build a red deer sperm bank by collecting and freezing
spermatozoa of the stags hunted in Hungary.
(Poster presentation.)
R 123
110
Immunohistochemical expression of androgen receptor (AR),
estrogen receptor alpha (ER) and estrogen receptor beta (ER) in
the caudal and metatarsal glands of sika deer (Cervus nippon).
M. Suzuki, Y. Yanagawa, Y. Matsuura, S. Otsuka, D. Hayakawa, M. Sasaki, C.
Yayota, H. Igota, S. Kondo, and N. Kitamura
Graduate School of Veterinary Medicine, Hokkaido University, Sapporo, Hokkaido,
Japan
Graduate School of Agriculture, Hokkaido University, Sapporo, Hokkaido, Japan
Department of Veterinary Sciences, Obihiro University of Agriculture and Veterinary
Medicine, Japan
The presence of hormone receptors has been reported in the skin appendages of cervid
species such as red and sika deer. According to previous studies, it has been indicated
that the mane follicles and skin glands of deer are under endocrinological regulations.
In this study, we investigated the immunohistochemical expression patterns of
androgen receptor (AR), estrogen receptor alpha (ERฮฑ) and estrogen receptor beta
(ERฮฒ) in the caudal and metatarsal glands of sika deer to determine whether these
large skin glands are regulated by sex steroid hormones. All samples were collected
from adult females and males shot for research purposes during the non-breeding
season (from March to May) on the island of Hokkaido, Japan. Immunoreactivities to
the three kinds of receptors were detected uniformly in the nuclei of secretory
epithelial cells of the caudal gland. These immunoreactivities were also observed in
cell nuclei of sebaceous glands and root sheaths of hair follicles in the metatarsal
gland. Expression patterns of these three receptors basically resembled each other in
both the caudal and metatarsal glands. In sebaceous glands of the metatarsal gland,
however, immunoreactivities of ERs tended to be weak and limited to cells near the
marginal area of the glands, as compared with those of AR. These results suggest that
secretions of the caudal and metatarsal glands and development of specific white hair
at the metatarsal gland area are influenced by sex steroid hormones in sika deer.
However, the difference in the steroid receptor immunoreactivities between females
and males was not clear in the present work. This may be due to the fact that all
samples were collected in the non-breeding season. Therefore, in order to fully discuss
sex differences in skin gland regulation systems, the same analysis should be
conducted by using samples obtained during the breeding season.
(Poster presentation.)
124 R
111
Comparison of estrogen receptor and progesterone receptor
expression during the estrus and pregnancy in uteri of sika deer
(Cervus nippon).
Y. Yanagawa
1
, Y. Matsuura
1
, D. Hayakawa
2
, C. Yayota
1
, M. Sasaki
2
, S. Kondo
1
, N.
Kitamura
2
, and M. Suzuki
1
1
Graduate School of Veterinary Medicine, Hokkaido University, N18-W9, Kita-ku,
Sappro, Hokkaido 060-0818, Japan
2
Obihiro University of Agriculture and Veterinary Medicine, Japan
Recently, the wild sika deer population has been increasing in Hokkaido, Japan, and
they cause serious problems such as agricultural damage and forest destruction. The
pregnancy rate of sika deer is very high, usually over 90% in a high quality
population. For the maintenance of pregnancy, it is necessary to inhibit luteolysis that
induced by PGF2รก secreted from the uterus. The PGF2รก secretion from uterus is
regulated by sex hormones; therefore, the expression pattern of receptors in the uterus
during pregnancy must be altered from that during the estrus. However, the
localization of hormone receptor in the uterus has not been reported in sika deer. In
this study, immunohistochemical expression of estrogen receptor (ER) and
progesterone receptor (PR) in the sika deer uterus during the estrus and pregnancy
period (early, middle, and late) were detected. Uteri were collected from wild sika
deer that were killed for damage control or research purpose. Localization and
immunoreactivities were detected visually in the luminal epithelium, gland, stroma,
and myometrium of uteri. During the estrus, both ER and PR were expressed in the
uterine cells such as shallower gland and stroma. As pregnancy progressed, the
expressions of ER and PR in the uterus become decreased and weaker than those of
estrus. In the late pregnancy period, both ER and PR could scarcely be detected, and
even when detected the immunoreactivities were very low. These results are
corresponded with previous reports on ewes wherein it was found that both the ER and
PR expressions were enhanced by estrogen and inhibited by progesterone. In the early
pregnant uterus of hinds, ER expressions were similar to estrus whereas those were
very weak even in very early pregnancy in cows and ewes.
(Poster presentation.)
R 125
112
Roaring trends in red deer: a preliminary analysis.
A. Bocci
1
, K. Attinault
2
, and M. Telford
1
1
Universitร di Siena, Dipartimento di Scienze Ambientali, Sezione di Ecologia
Comportamentale,
Etologia e Gestione della Fauna, via Mattioli 4 โ 53100 Siena, Italy
2
Ecole Nationale Superieure Agronomique de Rennes, 65 rue de Saint Brieuc 35042
Rennes, France
Red deer call loudly and repeatedly during the breeding season to assess dominance
between males (Clutton-Brock & Albon 1979). McKomb (1987) showed how roaring
can also induce/accelerate ovulation and that harem-holding males can improve their
mating success by calling repeatedly. Roaring rates change throughout the course of
the rut, with a slow increase of the number of roars/minute, a peak and a sharp decline
(Clutton-Brock & Albon 1979). We tried to study the roaring trend of a mature
population of red deer and its variability in relation to climatic variables. Within a
wider research project (red deer spatial behaviour, Eastern Italian Alps, Natural Park
of Paneveggio-Pale di San Martino), we studied some features of roaring behaviour
in 2004 and 2005. We noted down the number of roars/hour (roaring rate) during a
period of 24 hours (daily trend only for 2004) or 6 hours (trend during the whole
rutting period) to estimate the roaring trend. We started our observations 1 hour before
sunset, for the 6h period.
The 24h trend showed a peak 5-7 hours after sunset, with a significantly greater
roaring activity at night than in daylight (Mann-Whitney U test: n1=65, n2=55,
U=432, p=0.000). On the contrary, in Rhum Island, Clutton-Brock & Albon (1979)
found a diurnal consistent pattern, although with some variation across individuals.
As to the 6h period, there was a higher roaring intensity in 2004 (mean value for the
whole rutting period=143.9) than in 2005 (mean = 89.8); moreover, in 2004, the
roaring peak lasted longer (12 days) than in 2005 (9 days).
We estimated the influence of climatic variables on roaring (mean daily temperature,
precipitations and humidity). A significant negative correlation was found between
roaring rate and mean daily precipitation, in 2004, for the whole period (Spearman
rank correlation coefficient: N=15, R=-0.609, p=0.03) as well as during the peak
(Spearman rank correlation coefficient: N=9, R=-0.842, p=0.004). In both years, a
significant negative correlation was found between roaring rate and mean humidity
(Spearman rank correlation coefficient: 2004, N= 15, R=-0.736, p=0.02; 2005, N=13,
R=-0.582, p=0.04).
Acknowledgement: Parco Paneveggio-Pale di S. Martino, I. Angelini, E. Brunello, D.
Gabaglio, M. Ruggiero.
(Poster presentation.)
126 R
Behaviour and welfare
128 R
113
Assessing the performance of a Persian fallow deer population 10
years after reintroduction.
D. Saltz and S. Bar-David
Mitrani Dept. of Desert Ecology, Ben Gurion University and The Institute of
Evolution, Haifa University, Israel
We evaluated the size and range of a Persian fallow deer (Dama mesopotamica)
population in Israel, 10 year after its reintroduction began. We compared our results
with projections of a spatial-demographic model developed in 2002 based on data
from the first 3 years of the reintroduction. A total of 80 adult (>1 year old) females
were released between 1996-2005, of which 75 were radio-collared. We estimated
female population size using a modified Lincoln-Petersen index based on the
proportion of collared females amongst adult females sighted during 2005. We did this
in several stages. First, we estimated the number of collared females in the population
by simulating the survival for those females that were alive at the time their radio
failed and adding to it the number of females with transmitting radios at the end of
2005. Based on 5000 simulations, the mean number of collared females in the
population was estimated at 21.3 (95% CI =17-25). Second, based on direct sightings
during 2005 we estimated the proportion of collared females in the population as 0.29
with a S.E. 0.09 (N=24). Third, by multiplying each simulation output by a random
normal value generated from the estimated proportion and S.E. of collared females,
we constructed a distribution of population size estimates. The median of this
distribution was 73.3 and the low-end 95% CI was 46. The 2002 model predicted 67
females by the end of 10 years with a lower 95% CI of 54. Based on 66 direct
observations between 2003-2005 the population ranges over ~70 km
2
, which is also
in agreement with the 2002 model projections. The slow growth during the first 10
years was predicted by the model and is attributed to the low first-year post-release
survival (<85%) and reproductive success. The effect of these factors is expected to
subside as the ratio of captive- to wild-born animals declines. Thereafter, the
population is expected to grow exponentially and reach 400-600 females within 50
years. Observed growth of the wild Persian fallow deer population in Israel is
consistent with projections, and we conclude that it is performing well.
(Oral presentation.)
R 129
114
Social competence in Chinese muntjac deer.
A. Fischer and H. Hendrichs
University of Bielefeld, Dep. of Ethology, Morgenbreede 45, 33613 Bielefeld,
Germany
Subdominant animals in groups of mammals live under continuous strain due to the
presence of dominant individuals of the same sex. They are suppressed in different
ways and have to develop an ability to cope with this, using the space remaining in
their specific situations to generate well-being and to develop their species` specific
behavior. This situation of a subdominant animal is investigated in a morphologically
conservative deer species, the Chinese muntjac (Muntiacus reevesi micrurus), which
is mostly described as solitary living animal. Which spatial and social mechanisms
does it use and which abilities are required and developed while growing up and
remaining in a group ? Which internal states are involved and how are performances
individually shaped? In nearly 500 hours of observation breeding groups of three to
eight muntjacs of different ages, kept in enclosures of 1,800 - 2,500 m
2
, were studied,
recording about 250 behavioural units. The data concerning the use of the spatial
areas, different forms of activities, resting behaviour and social contacts, combined
with the accompanying postures indicating the internal states. With the arrangement
of a cartesian coordinate system it was possible to include the distance between two
animals at any time to estimate social relations influences. Discussed are individual
skills in coping with changing situations, the implications of social relationships and
the amount of experiences. First approaches to access the animals` knowlege about
their situation are tried. To describe and to analyse the different influences of the
various factors a model distinguishing three levels has been designed. This type of
study provides complementary insights into what constitutes a fully developed healthy
animal including its adequate motivational and affective states. Such knowledge is
important in different fields of research and application, for example it allows to
improve the animals' living conditions and to access in captive and free-living
individuals the extend of their actual well-being.
(Oral presentation.)
130 R
115
The analysis of sexual segregation in fallow deer (Dama dama) on
different time and space scales.
S. Ciuti, S. Luccarini, and M. Apollonio
Department of Zoology and Evolutionary Genetics, University of Sassari, Italy
Throughout the analysis of long-time series census data (1984-2003) and by means of
radio-telemetry (1997-2001; 23 females and 25 bucks), we assessed fallow deer spatial
and habitat sexual segregation on different spatial and temporal scales in the San
Rossore Estate (Italy). The combination of different studies performed in the same
area allows us to show that a detailed analysis can outline a situation in which more
than one theory is valid for explaining sexual segregation in the same species, as
probably several factors act on different temporal and spatial scales. From 1984 to
2003 females gradually abandoned the eastern disturbed sector of the Estate, the only
area affected by human presence (increased year by year from 1984), to the detriment
of the western undisturbed sector. Males remained in the eastern sector, in spite of the
increasing disturbance, supporting the predation risk hypothesis (large spatial and
large temporal scale). From 1989, males gradually increased their presence in the
disturbed sector, as they benefited from lower female density, avoiding areas with
higher female density, supporting therefore the indirect competition hypothesis (large
spatial and large temporal scale). Inter-sexual competition was pronounced in a small
area inside the undisturbed sector affected by habitat modification during the 1980s,
when a scrub area was converted to an open pasture, and this phenomenon further
incited males to leave this area and reach the disturbed sector, supporting again the
indirect competition hypothesis (small spatial and large temporal scale). As a final
result of this long term process, during the last five years we showed that males used
consistently disturbed areas, both during the day and the night; instead, females
frequented eastern disturbed areas only during the night, when human presence was
absent. Therefore, large scale segregation recorded during the day disappeared during
the night, when females usually reached the eastern sector, supporting again the
predation risk hypothesis (large spatial and small temporal scale). Nevertheless, sexes
segregated on a small scale inside this area, as they showed different habitat choices,
supporting the forage selection hypothesis (small spatial and small temporal scale).
(Oral presentation.)
R 131
116
Behavioural modifications of female ungulates during late
pregnancy and early lactation: the case of fallow deer Dama dama.
S. Grignolio, P. Bongi, S. Ciuti, E. Bertolotto, and M. Apollonio
Department of Zoology and Evolutionary Genetics, University of Sassari, Italy
Mammalian females exhibit complex behaviour patterns during weaning as a response
to energetic requirements and as antipredator behaviour, but this aspect is still widely
unknown for female ungulates. We analysed how pregnancy and lactation could
influence space use, habitat selection and time budget in female fallow deer. The study
was performed in two study areas (n = 23 and n = 15 adult females) of Tuscany (Italy)
using radio-tracking techniques and direct observation, respectively. Home range
analyses showed marked differential spatial behaviour between calving and non-
calving females only when fawns were present. Lactating does occupied smaller home
ranges than non-lactating ones only after the birth period. Habitat choices were
significantly different since late pregnancy, when mothers reached parturition site.
During parturition and lactation, calving females showed a higher use of marshes,
which provides good concealment for fawns. Calving females avoided rich areas with
short-grass meadows, since these are without any kind of concealment for fawns,
while they were selected by non-calving females because of their productivity. During
lactation mothers increased time spent foraging. Aimed to generalize behaviour
patterns of female ungulates during parturition season, we compared fallow deer
behaviour to those recorded in other ungulate researches. Furthermore, we made a
comparison with a study performed by our research group on Alpine ibex (Capra
ibex), a species phylogenetically distant (alpine ibex bovid vs. fallow deer cervid),
living on a different environment (Alpine habitat vs. Mediterranean habitat) and
characterized by a different young behaviour (follower strategy vs. hider strategy).
Even though these differences, ibex females (n = 28) showed a similar behaviour
during parturition season. Lactating ibex females showed smaller home range than
non-lactating ones after the birth period. Mothers preferred rocky slope areas and
likely they moved to sub-optimal but safer habitats during lactation in order to reduce
the predation risk for their offspring, and at the same time they increased food intake.
This study show that ungulate mothers may move to sub-optimal habitats,
compromising their energy intake, to reduce the predation risk to their neonates, but
compensating by spending more time foraging and having a higher bite rate.
(Oral presentation.)
132 R
117
Pre-orbital gland opening in red deer (Cervus elaphus) calves:
Signal of excitement?
J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, and L. ล vecovรก
Ethology Group, Research Institute of Animal Production, P.O.Box 1, Praha 10 -
Uhลรญnฤves, 104 01, Czech Republic
The opening of the pre-orbital gland of red deer (Cervus elaphus) calves has been
previously associated with feeding and achieving satiety. However, it has been
suggested to be most likely affected by some other factor or factors, possibly by
excitement of the calf. If so, a calf should open its pre-orbital gland while being
exposed to any stressful procedure. The hypothesis was tested that the pre-orbital
gland is closed in a relaxed calf while it is opened in a stressed calf. Pre-orbital
opening was observed in 41 newborn farm red deer calves during a regular daily
routine consisting of a search for new born calves, their inspection, weighing and
painful marking with an ear tag. The openness of the pre-orbital gland (pre-orbital
gland closed or opened) was recorded just prior to manipulation of a lying calf (i.e. in
a calm calf), and then during the manipulation (i.e. in a distressed calf). Prior to
manipulation, in all but three calves (7.32 %, all of which were males) the pre-orbital
gland was closed. All of the observed calves (100 %) opened their pre-orbital gland
during their manipulation, at least by the time when the ear was punctured by the ear
tag. Thirty eight calves (92.68 %) behaved along with the advanced hypothesis (
(1)
=
29.88, P < 0.0001). That is, the gland was closed prior to manipulation, and was open
after the calf was exposed to the stressful event. The openness of the pre-orbital gland
in newborn red deer calves was invariably associated with a stressful manipulation by
the humans. This suggests that it is likely a signal of calf excitement.
(Oral presentation.)
118
The effect of the birth weight on the calfโs allosucking success in the
red deer (Cervus elaphus) supports the compensation hypothesis.
A. Duลกek
1,2.
and L. Bartoลก
1
1
Ethology Group, Research Institute of Animal Production, Prague - Uhลรญnฤves, CZ-
104 01, Czech Republic
2
Biodiversity Research Group, Department of Zoology, Faculty of Science, Charles
University, Viniฤnรก St. 7, CZ-128 44 Prague, Czech Republic
The allosucking behaviour was studied in a number of ungulate species including the
red deer (Cervus elaphus). The allosucking calves are expected to be favoured over
R 133
the non-allosucking ones because of a surplus milk intake from non-maternal hinds.
In general, there are two reasons why the calves may be motivated to suck the non-
maternal hinds. The calves may try allosucking either (1) to gain competitive
advantage over their peers or (2) to minimize inter-individual differences in the body
size. The aim of this study was to test the second โcompensation hypothesisโ (Bartoลก
et al. 2001, Anim. Sci. 72: 493-500) presuming an increase in allosucking motivation
with the calfโs lower birth weight. Two groups of animals differing in the number of
hinds and calves (hinds/calves: n = 18/7 and 28/14) were observed. The (allo)sucking
success of the calf was expressed as a probability of its sucking by the hind. The
probability of successful sucking was modelled using logistic regression (proc
GENMOD for repeated measures, SAS V9) with predictors being the calfโs birth
weight, age, sex, fighting success and aggression of the allonursing hind tested. Out
of 1131 sucking events and 303 sucking attempts, we recorded 177 allosucking events
and 87 allosucking attempts, respectively. In agreement with the advanced hypothesis,
probability of the successful sucking decreased with the calfโs birth weight (ฯ
2(4)
=
10.38, p = 0.0345). This was affected by the calfโs sex and by the nursing status of the
hind. In terms of sucking, male calves were more successful than female calves
sucked by their mothers, while in non-maternal hinds it was the opposite. Probability
to suck decreased with the calfโs age (ฯ
2(1)
= 6.42, p = 0.0113). Relatively greater
allosucking success of the female calves might result from the inter-sexual difference
in the birth weight, with the female calves being significantly (t = 2.86, N
(M)
= 11, N
(F)
=10, p = 0.009) lighter than the male calves. Thus, our results support our
compensation hypothesis.
(Oral presentation.)
119
Do red deer (Cervus elaphus) grandmothers nurse their
grandchildren?
J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, J. Drรกbkovรก, L. ล vecovรก, J. Pluhรกฤek, R. Kotrba, A.
Duลกek
Ethology Group, Research Institute of Animal Production, POB 1, CZ-104 01 Prague-
Uhลรญnฤves, Czech Republic
Besides maternal investment to their progeny red deer hinds often nurse non-filial
offspring. However, causes of such behaviour are not yet fully understood. Across
studies, large individual variability in the incidence of allonursing has been reported.
It is presumed that the incidence of allonursing is influenced by kinship. Therefore,
we tested whether the hinds which nurse the non-filial calves are their grandmothers
or their matrilineal relatives. In three successive seasons, suckling behaviour was
observed in a herd of farmed red deer hinds having calves between parturition and
weaning. From 3952 sucking solicitations recorded in 57 calves mothered by 27 hinds
134 R
falling into 15 families, 1047 were immediately terminated by the female (โprevented
sucking attemptsโ) and 2905 were accepted (โnursingโ), 24.74 % of prevented sucking
attempts and 9.43 % of nursing being non-filial. Within non-filial sucking solicitations
when a lactating grandmother was present in the herd (13 calves, 240 solicitations),
all of the calves except one solicited its grandmother considerably less frequently
(10.00 % in total) than other non-maternal hinds (ฯ
2(1)
=16.76, P<0.0001; Fisher exact
probability test). The chance that a soliciting calf will be nursed (i.e. proportion of
nursing from all sucking solicitations for the calf) was 3.8-times higher in its
grandmother than in other non-maternal hinds. Nevertheless, this result disappeared
when adjusted for repeated measures within a calf, because the one calf predominantly
sucking its grandmother performed 83.33 % of all grandmothers solicitations
(ฯ
2(1)
=0.93, n.s.; logistic regression model, PROC GENMOD, SAS). In that
grandmother, the probability she will nurse her grandson did not differ from that of
her filial calf (0.90 vs. 0.86, respectively), which contrasted to significantly greater
probability of being nursed in filial (0.77) than non-filial (0.51) calves in general
(ฯ
2(1)
=11.21, P<0.001). We got similar results when we tested the effect of matrilineal
relatedness between a calf and allonursing hinds: few solicitations led to a relative
hind (12.77 % of 321), similar probability of being nursed in relative and non-relative
non-filial calves (0.46 vs. 0.47, n.s.), except of the reciprocal allonursing in
grandmother-daughter pair described above. Conclusion: The calves neither
predominantly solicit relative hinds nor the kinship between the hind and the
soliciting, non-filial calf would influence the hind decision to nurse it.
(Oral presentation.)
120
When prey fight back: higher levels of aggressive defence by mule
deer than whitetail females lowers vulnerability of mule deer fawns
to coyotes early in life.
S. Lingle
1, 2
, W. F. Wilson
1
, and S. M. Pellis
1
1
University of Lethbridge, Lethbridge AB T1K 3M4, Canada
2
University of Alberta, Edmonton AB T69 2E9, Canada
Female defence of young is widespread in ungulates, yet little work has been done to
investigate how this response affects predation on young. We observed naturally
occurring predatory encounters between coyotes (Canis latrans) and deer fawns to test
the hypothesis that a difference in aggressive defence leads to the differential
vulnerability of mule deer (Odocoileus hemionus) and white-tailed deer (O.
virginianus) fawns in summer, when fawns are 0 to 14 wks in age. Coyotes were less
likely to attack mule deer than whitetail fawns they encountered, and were less likely
to kill mule deer than whitetail fawns they attacked. The simple presence of a mule
deer female near a fawn deterred coyotes from attacking, and this was not the case for
R 135
whitetails. Once attacked, fawns of both species were less likely to be killed when
females defended them. However, mule deer females were far more likely to defend
fawns and banded together to defend fawns, resulting in fewer captures of mule deer
than whitetail fawns. These results indicate that higher levels of defence by mule deer
females contribute to the lower predation rates reported for mule deer than whitetail
fawns of this age group. It also raises the possibility that mule deer fawns, and even
whitetail fawns, may have improved survival during summer when occupying areas
with high numbers of mule deer females.
(Oral presentation.)
121
Why Help? The evolution of altruistic antipredator defence in mule
deer.
S. Lingle
1, 2
, D. Rendall
1
, and S. M. Pellis
1
1
University of Lethbridge, Lethbridge AB T1K 3M4, Canada
2
University of Alberta, Edmonton AB T69 2E9, Canada
Both white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus) females
defend fawns against coyotes (Canis latrans), but only mule deer defend non-
offspring conspecific and heterospecific fawns. At such times, females may have to
decide whether to defend a fawn having imperfect information on its identity obtained
from hearing a few distress calls. Playback experiments with fawn distress calls were
designed to consider the role of imperfect recognition while testing functional
hypotheses for fawn defence. Whitetail females only approached the speaker when
whitetail calls were played and when their offspring was hidden, suggesting fawn
defence was strictly a matter of parental care in this species. In contrast, mule deer
females responded similarly and strongly regardless of the callerโs identity (whitetail
versus mule deer; own offspring versus unfamiliar conspecific fawn), the femaleโs
reproductive state (mother or non-mother), even when accompanied by their own
offspring. This lack of behavioural discrimination revealed mule deer females did not
defend non-offspring because they have imperfect information on their own offspring.
Our results also suggest three traditional explanations for altruistic behaviour maternal
defence of the offspringโs area, kin selection and reciprocal altruism are unlikely to
account for the mule deerโs defence of non-offspring.
(Oral presentation.)
136 R
122
Cooperative anti-predatory behaviour in sympatric white-tailed,
fallow, roe and red deer: Experimental confirmation using a
dummy.
R. Kotrba, L. Bartoลก, J. Bartoลกovรก-Vรญchovรก, J. Panamรก, V. Kลกรกda, P. ล ustr, J.
Pluhรกฤek, A. Duลกek, D. Vaลkovรก-Formanovรก, G. Illmann, E. ล mรญdovรก, and K. V.
Miller
1
Ethology group, Research Institute of Animal Production, POB 1, CZ-104 01, Prague-
Uhลรญnฤves, Czech Republic;
1
Daniel B. Warnell School of Forest Resources, University of Georgia, Athens, GA
30602, USA
Our previous research has shown that grazing time in sympatric cervid species
increased if the focal deer was joined by another animal regardless of the species or
sex. We hypothesized that this served as an anti-predatory strategy based on
interspecific cooperative behaviour among the species (Bartoลก at al. 2002. J. Wildlife
Manage. 66, 522-527). To experimentally test our hypothesis, we used a plastic
dummy (โgrazing deerโ) with a remote control that could prone the dummy into
grazing position or lay the dummy down. From 2000 to 2005, we performed 52
observations (135 to 310 minutes each) in the Dobลรญลก Forest, Czech Republic on six
locations to get additional data for white-tailed deer (Odocoileus virginianus)- OV and
fallow deer (Dama dama)- DD, and to complete observations for additional two
species, red deer (Cervus elaphus)- CE and roe deer (Capreolus capreolus)- CC. We
used the dummy at one location, and simultaneously used one or two other locations
as controls. We switched between locations regularly and recorded the time spent on
the pasture for any deer present. On the location with the dummy, when any deer
entered the pasture and stayed there for 5 min, and was > 130 m from the dummy, we
erected the dummy and started the measurement. The analysis was performed by the
GLM model (SAS) with factors โDummyโ (with or without), โPresence or absence of
a dam with a fawnโ, โObserverโ, โLocationโ, โSize of a group entering the pastureโ
and continuous variables โTime until another deer entered the pastureโ and โNumber
of deer joining the focal animalโ. All species, OV, DD, CE and CC respectively, spent
more time with a dummy (LSMEANs ยฑ S.E.; OV 33.55 ยฑ 2.69 min, n= 49; DD 30.56
ยฑ 2.14 min, n= 50; CE 27.13 ยฑ 1.64 min, n= 45; CC 33.64 ยฑ 5.20, n= 13) than without
it (OV 21.22 ยฑ1.61, n=369, P<0.001; DD 24.01 ยฑ 1.00, n= 361, P<0.003; CE 16.80
ยฑ 1.42 min, n= 97; CC 23.41 ยฑ 2.81, n= 28). Thus, our experimental results supported
the hypothesis that all deer species, OV, DD, CE and CC, spent longer time on the
pasture if a dummy was erected than when it was not.
(Oral presentation.)
R 137
123
Rutting encounter between males and female choice in fallow deer
(Dama dama).
B. Friฤovรก
1
, L. Bartoลก
2
, J. Bartoลกovรก-Vรญchovรก
2
, J. Panamรก
2
, P. ล ustr
3
, and E. ล mรญdovรก
4
1
Department of Zoology, Faculty of Science, Charles University, Prague, Czech
Republic
2
Ethology Group, Research Institute of Animal Production, Prague, Czech Republic
3
Department of Science and Research, ล umava National Park Administration,
Kaลกperskรฉ Hory, Czech Republic
4
Department of Anthropology, Charles University, Prague, Czech Republic
In male fallow deer (Dama dama) conflicts and fights during the rut are a frequent
occurrence. It was postulated that a choice of the mate by a female is affected by the
results of these encounters. The hypothesis was tested that a female will stay in a
harem of the winner of an encounter, whereas she will abandon the loser. We observed
interactions between adult fallow bucks and the responses of fallow does in the
enclosure Bลezka (Czech Republic) during the rut in 1998-2000. For each encounter
between two opponents we recorded: a) which male initiated and which won the
encounter, b) did the encounter escalate into a physical fight, c) how many females
were present up to 20m from each of the males and d) where the females moved
during and after the encounter. A logistic regression model (PROC GENMOD, SAS)
was fitted to estimate the probability that a female will desert the male she had
accompanied before the encounter. Tested factors were: a) results of an encounter
(winner/loser/tie) for the male the female came with, b) whether the male was an
initiator of the encounter (yes/no), c) if the encounters escalated into a physical fight
(yes/no), d) the presence of females accompanying the opponent (yes/no). The
probability of a femaleโs desertion was affected by the outcome of an encounter
(ฯ
2(2)
= 7.44, p < 0.03) and presence of females by an opponent (ฯ
2(1)
= 10.15,
p < 0.01). Female desertion was more frequent in losers (0.26) than winners (0.07) or
in encounters without a clear outcome (0.16). When both opponents kept some
females, the probability of a female desertion was significantly lower (0.04) as
compared to the case when the male kept some females while his opponent had none
(0.41). Neither initiation of the encounter, nor escalation to physical fight affected the
probability of female desertion. Our data support the impact of an encounter outcome
on a decision of fallow does to stay with or leave a male she accompanied before the
encounter. The probability of a female to desert the male was generally low. That the
presence of females accompanying the opponent decreased the probability of female
desertion counteracts the mate-copy theory.
(Oral presentation.)
138 R
124
Habitat selection and home range size of red deer (Cervus elaphus)
in montane areas of ล umava National Park, Czech republic -
preliminary results.
P. ล ustr and A. Jirsa
Dept. of Science and Nature Protection, ล umava NP and PLA Administration, Suลกickรก
399, CZ-341 92 Kaลกperskรฉ Hory, Czech Republic
Home ranges of 10 male red deer Cervus elaphus (age 3-6 years) were studied in 2005
in National Park ล umava, Czech Republic using GPS telemetry. Each animal was
located 24 times a day. Winter enclosures are used for management of red deer in the
national park (NP) to prevent harms in surroundings of NP, thus only summer data are
available (May to November). Home range size ranged from 7,63 to 114,89 km
2
, mean
40,78 km
2
(MCP 95%). Two different behaviours may be distinguished - sedentary
(mean home range size 18,28 km
2
ยฑ 10,49) and migrant (68,90 km
2
ยฑ 32,57). Home
range size did not differ significantly during rut season in comparison to summer. NP
ล umava is covered mainly by spruce forest (more than 80%). Forest habitat was used
by red deer in 58,99 ยฑ 20,24% of locations, non-forested areas (meadows) was
selected preferably. There is a significant difference in habitat use during night and
day, non-forested areas are preferably selected by red deer during nighttime (high
level of anthropic disturbance). There is no significant difference in habitat use
between sedentary and migrant animals.
(Oral presentation.)
125
Sex-specific strategies of dentine depletion in red deer.
J. Carranza, C. Mateos, S. Alarcos, C. B. Sรกnchez-Prieto, and J. Valencia
Biology and Ethology Unit, University of Extremadura, 10071 Cรกceres, Spain
Worn teeth in herbivore ungulates may be related to lower efficiency in mastication
and hence lower performance. However, selection should favour maximal
performance in terms of body mass and reproductive capacity during reproductive
lifespan, when permanent teeth are already partially worn. We hypothesize that wear
rate may respond to a strategy of dentine expenditure, which balances instantaneous
wear rate and performance against dentine preservation for future performance and
reproduction. Here we study 4,208 carcasses of Iberian red deer Cervus elaphus
hispanicus to show not only that tooth wear rates were smaller in females than in
males but that they also showed different patterns of variation throughout lifetime:
while females showed a decelerated wear, males maintained a rather constant rate of
R 139
tooth wear. Most importantly, the relationship between tooth wear and body
performance also differed between the sexes: females with more worn teeth were
lighter but, in contrast, males with more worn teeth were heavier and had larger antlers
until senile age when more depleted teeth related to lower performance. These results
reveal for the first time sex-specific lifetime strategies of dentine expenditure:
maintenance of performance ability throughout a longer reproductive lifespan in
females, compared with maximizing performance by depleting dentine reserves within
a shorter lifespan in males.
(Oral presentation.)
126
Does a hindโs rank affect duration of filial and non-filial calfโs
nursing in red deer (Cervus elaphus)?
J. Drรกbkovรก, J. Bartoลกovรก-Vรญchovรก, L. Bartoลก, J. Pluhรกฤek, R. Kotrba, L. ล vecovรก, and
A. Duลกek
Ethology group, Research Institute of Animal Production, POB 1, CZ-104 01, Prague,
Czech Republic
The relationship between rank and suckling behaviour in red deer is controversial as
it has not been fully explained yet. . We presumed that the dominant position of the
hind yields profit associated with suckling behaviour for both the hind and her calf
(undisturbed longer suckling, no investment to non-filial calves etc.). We tested three
hypotheses: H1) The dominant hind should spend a longer time nursing than the
submissive one, H2) The dominant hind should not allonurse and H3) the dominant
hind should terminate nursing of her calf less frequently than the submissive one. Ad
libitum observation of suckling behaviour were made on a deer farm in a socially
stable group of 23 hinds and their 38 calves from birth to one month of the calfโs age
in two seasons. We used the Clutton-Brockโs fighting success index (โFSโ, Clutton-
Brock et al., Anim. Behav. 27:211-225) to determine a hindโs rank. Age of hinds
positively correlated with their rank. No change in the rank position occurred in the
two consecutive seasons (14 hinds were presented in both seasons). The impact of the
hind rank on suckling bout duration (H1) was tested using a generalized linear mixed
model approach (PROC MIXED, SAS, corrected for repeated measurements of
hind*calf pairs) with dependent variable โsuckling bout durationโ and fixed effects
โsexโ and โageโ of the calf, โFSโ nested within โseasonโ and interaction between โFSโ
and โsexโ. Suckling bout duration decreased with age of the calf (F
(1, 1592)
= 86.63 p <
0.0001). Neither of the remaining tested factors reached statistical significance. The
hindโs rank had no impact on suckling bout duration. The rank of the hind was not
correlated to her allonursing rate (H2; โFSโ*โnumber of allonursing/ all nursings in
one hindโ: r
s
= - 0.11, n.s.). The rank position had not explained the allonursing rate.
A logistic regression model (PROC GENMOD, SAS, corrected for repeated
measurements of hind*calf pairs) fitted to test H3 revealed that a probability of the
140 R
termination of the suckling bout by the hind increased with age of the calf (ฯ
2(1)
=
20.31, p < 0.001; odds ratio = 1.057), presumably due to weaning process. No effect
of hindโs rank was found either. Our data suggest that a hindโs rank does not affect
suckling behaviour in captive red deer.
(Poster presentation.)
127
ISAMUD: an integrated software environment for analysis and
management of GPS telemetry data.
F. Cagnacci
1
, F. Urbano
1
, C. Furlanello
2
, M. Neteler
2
, and L. Pedrotti
3
1
Centro di Ecologia Alpina, Viote del Monte Bondone (TN-Italy)
2
ITC-IRST, Trento (TN-Italy)
3
Consorzio del Parco Nazionale dello Stelvio Settore trentino Cogolo di Peio (TN-
Italy)
GPS technology has certainly represented a revolution for telemetry. Geographic
locations (XY coordinates) of collared animals can be obtained from satellites (via
VHF/GSM download or directly via ARGOS system), along with other data, such as
activity, outside temperature and precision of the localisation. However, the
potentially huge data sets collected by GPS collars may represent a challenge, both in
terms of data management and spatial analysis. 25 roe deer (16 females and 9 males)
have been captured and marked with GSM-GPS collars (Vectronic GPS-Pro 1D) from
January 2005 to date in Eastern Italian Alps (Trentino Province, Monte Bondone
area). Data are sent by the collars via GSM net, processed and downloaded as e-mail
messages and finally converted to dbf files. We developed an integrated software
environment to automatically store, associate geographic variables to and spatially
analyse the data. ISAMUD (Information System for Analysis and Management of
Ungulates Data) is mostly based on Open Source, Windows (Microsoft ) and Linux
operating softwares. The Geo-database PostgreSQL (http://www.postgresql.org/) and
its spatial geographic extension PostGIS (http://postgis.refractions.net) are the core of
the information system. Thanks to the latter, some geographic functions can be applied
directly in the database, without invoking an external GIS, e.g. some categorical
geographic variables (landuse, hunting districts) can be associated to the localisations.
Other more complex geographic features can be uploaded by the Open Source GIS
GRASS (Geographic Resources Analysis Support System, http://grass.itc.it/), now
available also in Windows version, via the interface Quantum GIS (QGIS)
(www.qgis.org). Statistical and geostatistical analyses, including Home Range
calculations, are carried out in R (http://www.r-project.org/), thanks to the numerous
available packages, Adehabitat among all (http://cran.r-
project.org/src/contrib/Descriptions/adehabitat.html), or to ad hoc developed
functions. Results are then stored in the main geodatabase, to be available for further
analyses. The database can be accessed by either PostgreSQL graphic interface,
R 141
including PGADMIN III (http://www.pgadmin.org/) and ACCESS
(www.microsoft.com/access/). The user-friendly data input/output format has been
developed within the latter. The flexibility of the system allows the development of
other modules and the applicability to different data sets from different species.
(Poster presentation.)
142 R
Censusing and modelling
populations
144 R
128
Censusing and modelling of red deer (Cervus elaphus L.)
populations in Poland by using โInventโ and โAntler-2000โ
software.
B. Bobek
1
, W. Frฤ
ckowiak
1
, M. Gawor
2
, M. Kolecki
1
, D. Merta
1
, and L. Wiลniowska
1
1
Department of Ecology, Wildlife Research and Ecoturism, Pedagogical University
of Cracov, Podbrzezie 3, 31-054 Krakow, Poland
2
Strata Mechanics Research Institute, Polish Academy of Sciences, Reymonta 27, 30-
059 Krakow, Poland
Computer program โInventโ was applied to estimate number of red deer inhabiting
155 Forest Districts (FD) covering 2.48 million ha of forest. In case of snow
conditions the input data include snow track density index (tracks/km) calculated on
basis of 5 days of tracking on line transects of 50 km per 10000 ha of forest area of
hunting district in each FD. It is also necessary to know relationship between
tracks/km and population density (N/1000 ha) which is established on basis of 500 ha
sampling plots. In case of lack of snow the input data include relative density index
i.e. number of animals seen per 1 km of strip transect (N/km) which were calculated
on basis of 3 days driving period and strip transects of 30 km per 10000 ha of forest.
Additionally relationship between N/km and N/1000 ha were calibrated on basis of
500 ha sampling plots and forest area of hunting district in each FD is required. The
obtained results indicated that population density between FD varied from 8.2-55.3
animals/1000 ha of forest.
The data on red deer number may be transported to โAntler-2000โ software that allows
to simulate number and structure of hunting bags under various hunting regimes.
Result of the simulations showed clearly that percent of calves in the total hunting bag
and percent of young males (2-5 years old) in the total harvested quotas of stags are
the most important variables influencing number of high trophy animals in the
population of red deer. Therefore percent of calves in the total hunting bag should not
be higher than 5%, while the percent of young males in the harvested quotas of stags
have to be maintained at the level of 30%. Practical application of โAntler-2000โ
program is presented and harvest strategy of the Polish red deer population is
discussed.
(Oral presentation.)
R 145
129
Estimating Red deer populations abundance in the Alps: successful
experiments on night surveys.
S. Focardi
1
, B. Franzetti
1
, A. Monaco
2
, and L. Pedrotti
3
1
Istituto Nazionale per la Fauna Selvatica. Via CaโFornacetta 9. 40064 Ozzano
Emilia (BO), Italy
2
Dipartimento di Scienze Ambientali. Universitร di Siena. Via P.A. Mattioli 4. 53100
Siena, Italy
3
Consorzio del Parco Nazionale dello Stelvio Settore Trentino. Via Roma 65. 38024
Cogolo di Peio (TN), Italy
The population size of red deer (Cervus elaphus) in the Alps has increased at a large
extent in the last decades, largely due to the presence of wide protected areas. The
Paneveggio-Pale di S. Martino Regional Park (PRP) and the Stelvio National Park
(SNP) (Trento province) show very high densities and red deer is becoming a relevant
problem. In order to undertake an effective management strategy, Parks require an
accurate assessment of actual population size and supported tests on both conventional
and innovative survey techniques. We present the results of a long-term project carried
out in the PRP and SNP devoted to compare distance sampling & thermal imaging
(DS & IR) with more conventional mark-resight (MR) surveys. Since 2003, in both
Parks, spotlight counts (SL) were undertaken in April-May (5-6 replicates/year) and
more then 20 radio-collared deer were available in each area to adjust survey results
for incomplete detectability using MR methods. DS & IR surveys were carried out in
parallel with SL & MR counts. A total of 20 and 45 kilometres of forest roads and
mountain tracks were walked at PRP and SNP, respectively. Each survey involved 2
observers and lasted 3-6 nights/area. Transects were surveyed between 10 p.m. and
5 a.m. The infrared camera was equipped with a laser range finder and an electronic
compass to measure observer-deer distance and angle without substantial error. Mean
population size ranges from 700 deer in the PRP to 1600 deer in the SNP. SL & MR
and DS & IR surveys provide coherent estimates (< 10% of relative mean difference),
both characterised by reasonably low CVs (< 30%). The results are very encouraging
and the proposed methodology may be potentially useful in many situations where an
accurate population estimate has to be collected. Survey design and costs are
described and discuss.
(Oral presentation.)
146 R
130
Whitetailed Deer Density Estimation Using Thermal Infrared
Imaging.
P. A. Tappe and R. E. Kissell, Jr.
School of Forest Resources, Arkansas Forest Resources Center, University of
Arkansas at Monticello, Monticello, Arkansas 71656, USA
The thermal infrared portion of the electromagnetic spectrum was discovered in
the latter part of the 17th century, with little research conducted on the uses of thermal
infrared imaging (TIR) until the 1920s. Croon et al. (1968) were the first to describe
the application of TIR for counting white-tailed deer (Odocoileus virginianus) using
a thermal imager mounted to a fixed-wing aircraft. Since that time, TIR technology
has continued to advance. Over the past decade, interest has increased in applying TIR
to estimate white-tailed deer densities, primarily because of questions and concerns
surrounding the precision and accuracy of other estimation methods. However,
inconsistency in sampling design and analysis has produced mixed results. Thus, we
initiated studies to investigate the use of ground-based and aerial TIR for estimating
white-tailed deer density in eastern Arkansas. Specifically, we compared density
estimates between ground-based TIR and spotlight counts (SLC) using 2 estimators,
and evaluated detection rates and compared 3 density estimators using aerial TIR.
Studies took place on 5 sites in eastern Arkansas within the Mississippi alluvial
plain: White River National Wildlife Refuge (WRNWR), Choctaw Island Wildlife
Management Area (CIWMA), Cut-Off Creek Wildlife Management Area (CCWMA),
Wingmead Farms (WMF), and Lakeside Hunting Club (LHC). The WRNWR
encompassed 87 km of the lower White River and was composed primarily of
bottomland hardwood forests (85%) with the remainder (15%) being sloughs, oxbow
lakes, and agriculture lands. Topography was flat and flooding occurred annually. All
other sites were composed of varying proportions of bottomland hardwoods and
agricultural fields. Topography was flat and the areas were subject to seasonal (winter-
spring) flooding.
A Mitsubishi IR โ M700 thermal infrared imager was used for all studies. The
imager had a sensor array of 801 x 512 pixels, a sensitivity of < 0.08ยฐC, and used a
50 mm, f1.2 silicon lens (14
o
[h] x 11
o
[v] field of view). It detected electromagnetic
wavelengths from 1.2 to 5.9 ยตm, which consisted of reflective infrared (1.2 to 2.9 ยตm)
and short wave thermal infrared (3 to 5.9 ยตm). Output was conducted through an
RS170, 75 โฆ connection to a GV-D1000 MiniDV Video Walkman digital video
recorder (Sony, New York City, NY).
For aerial surveys, we used a Cessna 182 fixed-winged aircraft with the imager
mounted in the belly of the plane. Surveys were flown at 457 m above ground level
with transects 110 m wide and at a speed of 130 kph. A global positioning system
(GPS) signal was routed through a video encoder-decoder (VED), and locations of the
plane obtained from the GPS unit were recorded on the audio portion of the video
tape. The VED labeled the video with a continuous stream of positions as well as time,
date, speed, and altitude information. GPS locations, obtained once each second, were
R 147
used to geo-reference frames on the digital tape. Double counting was prevented by
use of GPS locations integrated with videography. For ground-based surveys, we
mounted the imager on a tripod placed in the back of a pickup truck and recorded
digital video as described above. All video was reviewed using a high resolution, 1000
line, black and white Sony PVM-137 33-cm monitor.
In 2002 โ 2003 on WRNWR, we compared densities derived using 2 methods,
SLC and ground-based TIR, using both fixed area (FA) and distance sampling (DS)
estimators. Three survey routes (portions of existing roads), North Unit (10.0 km),
Jackโs Bay (12.7 km), and Ethel (24.6 km), were sampled in winter (December โ
March). For each survey route, SLC and ground-based TIR were conducted 6 โ 7
times each. A hand-held, 700,000 candle-power spotlight was used for SLC. All
surveys commenced 1 hr after sunset. The area visible from the transect was measured
perpendicularly on each route using a laser rangefinder at 0.16 km intervals in order
to compute area for the FA density estimate (number of deer sighted / unit area).
Program DISTANCE was used to estimate deer densities using DS (Buckland et al.,
1993) for each route. Data from repeated surveys were combined for each route to
increase sample sizes for both TIR and SLC. Appropriate models were chosen using
ฯ
2
goodness-of-fit (ฮฑ = 0.05) and lowest Akaike information criteria values. TIR
detected 3.4 times more deer and 2.4 times more groups of deer than SLC (Table 1).
Table 1. Mean numbers (SE) of individuals and groups of white-tailed deer detected
using a spotlight and ground-based thermal infrared imaging.
Variable Route Method
Spotlight Thermal
No. of Deer North Unit 1.4 (0.97) 9.9 (2.96)
Jackโs Bay 9.8 (2.33) 22.2 (4.36)
Ethel 6.0 (1.55) 6.8 (2.04)
No. of Groups North Unit 1.0 (0.58) 4.1 (1.12)
Jackโs Bay 4.8 (1.01) 9.5 (2.11)
Ethel 3.2 (0.87) 3.8 (0.87)
Densities derived from each combination of method and estimator could be ranked
in decreasing order as TIR
FA
= TIR
DS
>= SLC
FA
= SLC
DS
(Table 2). However, the
coefficient of variation (CV) was high for all estimators ( >= 25%). Though we did
not know the true densities, we suggest TIR combined with FA or DS produced the
least biased density estimates because deer/ha computed using FA represents a
minimum density.
148 R
Table 2. Mean densities (deer/ha) and SE of white-tailed deer obtained from spotlight
counts and thermal infrared imaging using fixed area and distance sampling
estimators.
Route Method Density SE
North Unit
TIR
FA
0.16 0.05
TIR
DS
0.14 0.03
SLC
FA
0.02 0.02
SLC
DS
0.03 0.02
Jackโs Bay
TIR
FA
0.46 0.09
TIR
DS
0.25 0.06
SLC
FA
0.20 0.05
SLC
DS
0.13 0.04
Ethel
TIR
FA
0.06 0.02
TIR
DS
0.03 0.08
SLC
FA
0.05 0.01
SLC
DS
0.03 0.01
In March 2003, we assessed the aerial detection rate of white-tailed deer on
CIWMA using human surrogates. Body temperatures of deer range from
approximately 37.2 ยฐC to 39.4 ยฐC (DelGuidice et al., 2001), whereas the average
human body temperature is 37ยฐC. Preliminary work indicated humans have slightly
lower thermal signatures than deer (Kissell, unpub. data); therefore, slightly lower
detection rates were expected. We substituted humans for deer to provide surrogate
minimum detection rates. Given humans have lower thermal signatures than deer, we
assumed that if we could detect humans we would be able to detect deer. Individual
persons (n = 20) were sent to random, known locations within the study site.
Individuals assumed a reclined or horizontal position on the ground to simulate the
dorsal surface area of a deer. Four of the 20 people were located in water and 16 on
dry land. Slightly overlapping, parallel transects were flown between 2000 hrs and
2200 hrs. The maximum temperature during the day preceding the flight was 18.3 ยฐC,
and the temperature declined from 12.2 ยฐC to 10.6 ยฐC during the flight. Thermal
signatures were recorded as people, deer, possible deer, possible people, or unknown.
Known locations were compared to locations identified from the video and the
percentage of people correctly identified was calculated. We detected 75.0% of the
people across the area and 93.8% of the people when the effect of water was taken
R 149
into consideration (Table 3). Thermal signatures of people and deer occupying flooded
areas were masked by the surrounding thermal signature of water. We found the
detection rate was very high in bottomland hardwood forests under dry conditions.
Table 3. Number of thermal signatures classified as people or deer using aerial thermal
infrared imaging.
Detected Identified As
People Deer
People 15 10 5
Deer 55 0 55
Following our investigation of detection rates, we used aerial TIR to compare deer
densities based on FA, mark-resight (MR), and DS estimates. Each of 4 study sites
(CIWMA, CCWMA, WMF, and LHC) was surveyed 4 nights each in February 2004.
Non-overlapping, parallel transects were flown 400 m apart between 2300 and 0600
hrs. FA densities for each night were calculated, using transects as replicates, as the
number of deer/unit area, and then averaged over 4 nights to derive a mean density for
each study site. MR densities were determined by using 2 independent observers to
review video for each night and record the time and location of each thermal signature.
Deer detected by the first observer represented the marked population, and deer
identified by the second observer represented the resighted population. Program
NOREMARK (White, 1996) was used to derive a density using the joint
hypergeometric maximum likelihood estimator, with transects as replicates, for each
study site. DS densities were calculated by delineating transects on video images,
measuring the perpendicular distance to each deer detected, and entering the
information into program DISTANCE. Appropriate models were chosen using ฯ
2
goodness-of-fit (ฮฑ = 0.05) and lowest Akaike information criteria values. Densities
derived from each estimator could be ranked in decreasing order as MR >= FA = DS
(Table 4).
Table 4. Mean densities (deer/ha) and SE of white-tailed deer obtained from aerial
thermal infrared imaging using fixed area, distance sampling, and mark-resight
estimators.
Location Method Density SE
CIWMA
FA 0.26 0.01
DS 0.26 0.02
MR 0.32 0.00
CCWMA FA 0.11 0.06
150 R
Location Method Density SE
DS 0.10 0.02
MR 0.18 0.00
LHC
FA 0.06 0.16
DS 0.06 0.01
MR 0.09 0.00
WMF
FA 0.20 0.02
DS 0.23 0.05
MR 0.37 0.00
Though we did not know the true densities, we suggest MR produced the least
biased estimates. The lower bounds of the MR estimates were based on the actually
numbers of deer observed, and the numbers of deer observed using MR were greater
than those observed using either FA or DS. This difference was likely due to 2
observers independently reviewing each tape for MR estimates, as opposed to 1
observer for FA and DS estimates. Regardless, MR density estimates were more
precise (CV = 0.0%) than FA (CV = 18.6 โ 46.1%) or DS (CV = 8.8 โ 26.9%)
estimators because of the large number of marked and resighted animals.
Though interest in the use of TIR has increased in the last decade, there are few
studies that have addressed its potential for estimating density of white-tailed deer. We
have found that ground-based and aerial TIR have great potential for increasing the
detection of deer and enhancing our ability to estimate deer densities. However,
research using a known population to investigate accuracy of estimators using TIR
data is needed. Additionally, all of our investigations have been concentrated in
bottomland hardwood ecosystems of the southern United States. Further investigations
are warranted to assess the performance of TIR in other geographic and environmental
conditions.
References
Buckland, S. T., Anderson, D. R., Burnham, K. P., Laake, J. L. (1993) Distance
sampling: estimating abundance of biological populations. Chapman and Hall,
London.
Croon, G. W., McCullough, D. R., Olson, C. E., Queal, L. M. (1968) Infrared
scanning techniques for big game censusing. Journal of Wildlife Management 32:751-
760.
DelGuidice, G. D., Mangipane, B. A., Sampson, B. A., Kochanny, C. O. (2001)
Chemical immobilization, body temperature, and post-release mortality of white-tailed
R 151
deer captured by clover trap and net-gun. Wildlife Society Bulletin 29:1147-1157.
White, G.C. (1996) NOREMARK: Population estimation from mark-resight surveys.
Wildlife Society Bulletin 24:50-52.
(Oral presentation.)
131
Estimating red deer Cervus elaphus populations: an analysis of
variation and cost effectiveness of counting methods.
M. J. Daniels
Deer Commission for Scotland, 82 Fairfield Road, Inverness IV3 5LH, UK
Different counting methods are currently used to estimate red deer populations in the
open range in Scotland but there are few data available to compare variation in
estimates, or relative cost-effectiveness. While it is impossible to determine the
accuracy of counts (as real numbers are unknown), variation within and between
different methods can be measured by repeat counts of the same area within a short
period as possible. This study aimed to quantify the variation observed from repeat
counts by each of four methods (ground, helicopter, infra-red helicopter and dung
counting methods) at one of three study sites in late winters 2003, 2004 and 2005.
Additional data from digital camera images of groups from counts in other areas of
Scotland were also used to assess the accuracy of visual counts. Coefficients of
variation within any method of between 5% and 16% were recorded, consistent with
previous comparisons for red deer open range counts in Scotland. Coefficients of
variation were lowest for ground and helicopter counts. The infrequency of optimal
conditions was likely to limit the applicability of infra-red counts in Scotland. In terms
of cost-effectiveness, helicopter counting was the least labour-intensive, with costs of
other techniques depending on the availability of existing manpower as an overhead
cost. It is concluded that helicopter counts are most likely to minimise errors while
maximising cost-efficiency. Accuracy can be improved by the use of digital
photography for counting larger deer groups. Estimates are likely to be improved
further by increasing the frequency of counts and using the same methods, counters
and routes for repeat counts.
(Oral presentation.)
152 R
132
Simple Movement Models for Complex Animals in Heterogeneous
Landscapes.
J. M. Morales
University of Cambridge, and Universidad Nacional del Comahue, Statslab CMS,
Wilberforce Road, Cambridge CB3 0WB, UK
Most animals have important cognitive capacities which are evident in their movement
trajectories. In particular, being able to remember locations and to return to them
contrast with simple movement models such as Fickian Diffusion or correlated
random walks. I present movement model formulations based on the premises that the
complexities of animal movement can be dissected into a few general movement
strategies, and that individuals modulate the switch among these strategies as they are
affected by changes in the internal and external environment. The models are
parameterized with North American elk GPS-tracking data using Markov Chain
Monte Carlo methods.
(Oral presentation.)
133
Reconstruction of the male population of red deer in Hungary.
S. Csรกnyi
Department of Wildlife Biology and Management, Szent Istvรกn University, H-2103
Gรถdรถllรต, Hungary
The reliability of spring population reports and the size of red deer (Cervus elaphus)
population had been debated recurrently during the last decades. Age data collected
on the antlers of stags presented for trophy evaluation provides an opportunity for
retrospective analysis of population size and trends. For this purpose I collected the
data published by the National Trophy Evaluation Committee from 1973 to 2005.
Based on these data I developed a simple population reconstruction model to calculate
the male cohort sizes and the population sizes for the period of 1973 and 1995. I
compared the reconstructed male population sizes with the reported male numbers in
the annual game management statistics. Marked differences have been found between
reported and reconstructed numbers, e.g., the reports seriously understated the stag
population. In this case population reconstruction gives a unique opportunity to reveal
the bias of reported numbers, and to study the survival pattern of males in harvested
populations. Although, the model is a deterministic one and it can only be used to
calculate population sizes in the past, its results contribute to the understanding of
R 153
population dynamics and the effects of harvest.
(Oral presentation.)
134
The second mass-mortality of an introduced sika deer population.
H. Takahashi and K. Kaji
Forestry and Forest Products Research Institute, Tokyo University of Agriculture and
Technology, Momoyama, Kyoto, 612-0855, Japan
Mass-mortality of ungulate populations sometimes occurred due to starvation after
they degraded vegetation. To clarify how an ungulate population behaves under food
limitation after the mass-mortality, we monitored population size, resource use and
mortality of a sika deer (Cervus nippon) population for 13 years from 1992 to 2004.
On Nakanoshima Island inside in Lake Toya, Hokkaido, northern Japan, the sika deer
population originated in three introduced individuals between 1956 and 1965. The
population irrupted up to 299 deer in 1983. The first mass-mortality occurred in 1984
following disappearance of food plants including dwarf bamboos, Sasa spp. At least
67 deer died, and 95 deer were translocated outside the island. Then the population
size was approximately halved. Thereafter the deer shifted their main food into fallen
leaves of deciduous broad-leaved trees throughout years. In addition, the deer
gradually increased use of unpalatable plants such as plum-yew, Cephalotaxus
harringtonia var. nana, which had expanded to vacant spaces after disappearance of
dwarf bamboos. The sika population re-increased over 400 deer in 2001, and
availability of plum-yew decreased to nearly zero until 2003. Then, the second mass-
mortality occurred (at least 113 deer died) in 2004 irrespective of milder winter than
usual. Annual and winter mortality rates were lower than females particularly during
the mass-mortality. Higher tolerance to habitat degradation may cause lower mortality
in females than in males. This could allow the population to increase after the first
mass-mortality where the vegetation has not recovered.
(Oral presentation.)
154 R
135
Fecal-pellet group count as index of sika-deer (Cervus nippon)
population density on subalpine plateau in Japan.
R. Goda, M. Ando, H. Sato, and E. Shibata
Graduate School of Bioagricultural Sciences, Nagoya University, Nagoya 464-8601,
Japan
Kyoto Prefectural Nantan Regional Promotion Office, Kyoto621-0851, Japan
Faculty of Science, Nara Womenโs University, Nara 630-8506, Japan
To assess the validity of fecal-pellet group counts as an index of sika-deer (Cervus
nippon) population density, we examined the relationship between deer density
estimated by a block-count method and fecal-pellet group counts on the Ohdaigahara
Subalpine Plateau in central Japan in spring (April 2006), summer (July 2005) and
autumn (October 2005). The study area (567 ha) was divided into six blocks according
to vegetation and topography. Each block was further divided into several sub-blocks
bounded by streams and ridges. As we walked the sub-blocks, we observed and
recorded details of sika deer, such as sex and age class, including the time of
observation and direction of movement. Furthermore, we set up a transect (1 m - 100
m) in each sub-block, counting fecal-pellet groups (>10 fecal pellets) within it. As a
result, the overall estimated deer population density in spring was 12.3/km
2
, ranging
from 6.0 to 54.0/km
2
. In summer, it was 17.3/km
2
, ranging from 0 to 43.5/km
2
and in
autumn, it was 12.3/km
2
, ranging from 9.6 to 33.1/km
2
. Fecal-pellet group counts
showed a significant positive correlation with deer density in spring (r = 0.81) and
autumn (r = 0.88). No significant correlation was found in summer, probably because
sika deer concentrate in the grasslands of Sasa nipponica, which is their main summer
forage and is high in nutrients. Therefore, we conclude that fecal-pellet groups should
be counted in spring or autumn to give a valid index of population density.
(Poster presentation.)
136
Comparison of four techniques to estimate roe deer abundance in
Alpine areas.
N. Putzu, V. La Morgia, and F. Bona
Universitร degli Studi di Torino, Dipartimento di Biologia Animale
via Accademia Albertina 13 - 10123 Torino, Italy
This research was carried out in 2005 in Chisone, Pellice and Germanasca Valleys,
Western Italian Alps (Piedmont, Italy). The aim of the study was to compare roe deer
density estimates obtained by different survey methods: pellet count (clearance plot
R 155
sampling and standing crop strip transect counts), distance sampling and block census
techniques were applied in sample areas, while data for the whole study area were
obtained both through distance sampling and block census methods. Simulations were
designed to study the performance of the methods under a wide variety of conditions.
Locations of animals were generated (1) according to a pseudorandom uniform
distribution and (2) according to presence probabilities derived from habitat evaluation
procedures. In simulations, roe deer density was set to 1, 3, 6, 9 or 12 animals/km
2
;
survey area ranged from 0.01% to 11% (5x5 m or 10x10 m plots). Our results
indicated that in sample areas density estimates obtained by pellet counts with faecal
accumulation rates were quite similar to those obtained by block census, while
standing crop strip transect counts underestimated the number of animals, maybe
because estimates were strongly dependent on accumulation rates, which we derived
from literature data. Anyway, plot sampling simulations indicated that a great effort
was needed to reduce coefficient of variation and to obtain narrow confidence
intervals of estimates.
Analysis of distance sampling data was complicated by the limited number of sighted
groups during the surveys at both scales of analysis. This technique slightly
underestimated roe deer population even in the whole study area, but a limited
increase in sampling effort may strongly improve the performance of the method.
We then suggest the use of distance sampling to estimate roe deer abundance in
Alpine areas, as pellet counts require a bigger sampling effort and may be unsuitable
for assessing roe deer abundance in large study areas; moreover, in our study pellet
counts were seriously affected by uncorrect estimation of accumulation rates.
(Poster presentation.)
137
Distance sampling and pellet group count to assess deer
populations: an application to conservation and management in the
Alps.
L. Pedrotti
1
, F. Cagnacci
2
, I. Callovi
1
and A. Tagliabรฒ
2
1
Consorzio del Parco Nazionale dello Stelvio via Roma 65, 38024 Cogolo di Peio
(TN-Italy)
2
Centro di Ecologia Alpina, 38040 Viote del Monte Bondone (TN-Italy)
In Italian Eastern Alps, deer (i.e. red deer and roe deer) have undergone contrasting
changes in density and geographic distribution in the last 10 years. This context may
require management/conservation actions for aiming at a balanced deer population
structure. Censusing methods commonly applied in game management (e.g. drives,
block censuses) may not prove robust and precise enough to allow for short term
validation. Pellet group count with distance sampling seems to encompass
randomness, replicability, comparability of results and possibility to aim at a certain
156 R
level of precision controlling the effort. In this paper, we assessed the feasibility and
efficacy of this method in two areas of the Alpine range: red deer density was
estimated in Stelvio National Park - Trentino sector (P.N.S.), whilst roe deer density
was estimated in Monte Bondone area - Trento Province (M.B.). In both areas,
ongoing telemetry studies allowed mark-resight estimates, considered as reference
density values. Systematic grids of line transects were randomly over imposed to the
study areas and sampled both in spring (winter density) and autumn (summer density),
for a total survey length of 24 km (n=240 transects, 100 m each) in P.N.S. and 23 km
(n= 115 transects, 200 m each) in M.B. Meanwhile, a retrospective decay rate
experiment was carried out in both areas: fresh pellet groups were placed monthly in
different habitat conditions and later checked for their persistence. A total of 9792 red
deer pellet groups were counted in PNS, corresponding to a highly precise pellet
groups density value (winter: d= 1779.2/ha, c.v.= 7.1%; summer: d= 1740.9/ha, c.v.=
11.6%). A total of 481 roe deer pellet groups were counted in M.B., leading to a less
precise, but close to the target, density value (winter: d= 123.2/ha, c.v.= 19.6%;
summer: d= 107.9/ha, c.v.= 18.3%). Density estimates proved to be robust to
environmental factors such as habitat type, exposition, slope. However, precision
decreased when the variability due to the estimated decay rate was taken into account
to calculate deer density from pellet groups density. The range of density values was
comparable with mark-resight results. Pellet group count with distance sampling
proved a robust method for deer population assessment in wooded areas and difficult
Alpine terrain, both with high and low densities, either concentrated or even
distribution of the populations. However, if pellet group count with distance sampling
were routinely used for deer population density estimate, a continuous monitoring of
decay rate would be strongly suggested. Management efficacy in terms of costs and
effort required is also discussed in comparison with other methods.
(Poster presentation.)
138
Red deer (Cervus elaphus) space use and population dynamics in
two Alpine National Parks.
F. Filli, L. Pedrotti, and H. Gunsch
Parc Naziunal Svizzer, CH-7530 Zernez, Switzerland
Stelvio National Park, Via Roma 65 - 38024 Cogolo di Peio (Trento, Italy)
Stelvio National Park, Piazza Municipio 1 - 39024 Glurns (Bolzano, Italy)
Having faced almost complete extinction in the Central Alps, red deer (Cervus
elaphus) experienced a real come back at the beginning of the 20th century. Since
then, their populations have been strongly increasing both in the Swiss National Park
(SNP) and the Stilfser Joch National Park (SJNP). Their typical migrations between
summer and winter ranges and their great adaptability pose difficult tasks for the
hunting authorities as well as the people responsible for the protected areas.
Population trend was investigated through different abundance estimates from 1996
R 157
to 2006 in the management unit including Trentino sector of SJNP and surrounding
hunting preserves of Val di Sole (720 km
2
) and cohort analysis allowed the
reconstruction of population dynamic from 1973. Red deer reached its carrying
capacity in 1999 when natural mortality started to significantly contribute to
population regulation. Contrary to Swiss experience, hunting activities could not
regulate population growth, as population progressively gathered in wintering areas
inside the park. 21 female red deer were equipped with transmitters in the SNP and 35
female red deer in the SJNP within the framework of a co-ordinated project. Their
home ranges were determined using one of the classical kernel method. In the SJNP
their summer home range amounts to 443.3 ha in the South Tyrol sector and to 960.1
ha in the Trentino sector and their winter home ranges to 384 ha and 501 ha,
respectively. In the SNP their summer home range amounts to 486.9 ha in the Il Fuorn
area and to 324.4 ha in the Val Trupchun area and their winter home ranges to 586.7
ha and 1391.8 ha, respectively. To migrate between winter and summer ranges the
animals cover distances of up to 16 km in the SNP area compared to 4.5 km in the
SJNP. We assume that the quality of the home ranges determines the home range
sizes, the lack of disturbances being the most essential resource both in the SNP and
SJNP. Despite similar population density in the different study areas, condition (body
mass) and constitution (jaw and hind foot length) index showed differences, according
to habitat quality and climate.
(Poster presentation.)
139
A population-dynamic study of red deer in Baranya, Somogy, Tolna
and Zala counties from 1970 to 2006.
R. Barna and L. Sugรกr
University of Kaposvรกr, Faculty of Economic Science & Faculty of Animal Science,
Hungary
The quality of the Southern Trans Danubian regions red deer stock is well above the
world average. This is verified by the fact that from the 17 Hungarian red deer
trophies listed among the worlds Top 50, 13 comes from the region studied.
With the aid of the created population dynamic age group model, based on the cover
data available form 1970, we have calculated the red deer populations in Baranya,
Somogy, Tolna and Zala Counties. Based on this model there were 13.678 red deer
in the four counties altogether. That is more than double of the amount estimated by
game managers. By 1990 the population raised to 49.298. After reducing the
population in 1993 it fell to 47.122. Then by 2000 it climbed to the maximum of
62.278. Because the numbers have been being strongly reduced since then, today there
are as few as 25.277 animals. Calculations based on the model show that the original
1:1.5 sex ratio has changed radically as a result of the forced killing of females. Today
there are more stags than females. This fact is supported by the field observation
158 R
results too. There are about 30.484 trophy data accessible since 1974 in the Somogy
County Trophy Evaluation Database. By analyzing these data a study was conducted
on the changing quality of the population. In age group 4 to 11 years the average mass
of antlers increased in 2004 and reached a 32 years period maximum in 2005. This
means that the previously saved stronger stags are killed now even at a young age.
Because of this the ratio of the older stags and so the medal awarding trophies are
reduced. The red deer population is becoming too young. In 2005 the penalties and
monetary sanctions were revoked. The amount of the so called negative mark trophies
is 8 times (142:17) higher than the 2004 value.
(Poster presentation.)
Antler biology
160 R
140
Visualization and characterization of stem cells from the
regenerating deer antler.
H. J. Rolf
1
, U. Kierdorf
2
, H. Kierdorf
2
, N. Seymour
1
, J. Napp
1
, H. Schliephake
1
, and
K. G. Wiese
1
1
University Hospital, Department of Oral and Maxillofacial Surgery, Clinical
Research, University of Goettingen, D-37075 Goettingen, Germany
2
Department of Biology, University of Hildesheim, D-31141 Hildesheim, Germany
It has recently been suggested by different authors that antler regeneration is a stem
cell based process; however, the presence of stem cells in the regenerating antler has
thus far not been demonstrated. The aim of the present study was therefore to
investigate whether cells positive for known stem cell markers are present in the
regenerating antlers of fallow deer (Dama dama). Using immunostaining, cells
positive for the mesenchymal stem cell markers STRO-1 and CD133 were, for the first
time, detected in biopsies taken from the cartilaginous tip of the regenerating antler.
Here, Stro-1+ cells were found mainly as small groups around vessels and capillaries,
thus representing perivascular cells. By cell sorting (FACS and MACS
ยฎ
up to 4%
STRO-1+ cells and up to 11% CD133+ cells could be obtained from primary mixed
cell cultures derived from the biopsies. To examine their developmental plasticity in
vitro, we tested the cells displaying the stem cell markers under different culture
conditions. Over 90% of the purified STRO-1+ cells remained positive after four
weeks of cultivation; subsequently progressive differentiation caused a decline in
number of STRO-1+ cells in the culture to only 0.4% after 12 weeks. In contrast to
mixed primary antler cell cultures, which contain different cell populations and are
able to form complex, three-dimensional structures in vitro, structure forming capacity
of antler stem cells seems to be restricted. Depending on the culture medium, Stro-1+
cells formed small nodules and stem cell typical ''cobblestone areas'', while CD133+
cells mainly formed nodule-like colonies. When the cells were cultured in a specific
neuroblast medium containing NGF, completely different structures were formed. Our
results suggest that the cartilaginous tip of a regenerating antler contains a vascular-
associated pool of adult stem cells with a high degree of phenotypic plasticity. This
makes the deer antler a useful model for the study of the role of stem cells in organ
regeneration in mammals.
This study was supported by the DFG (grant no RO 2520/1-1).
(Oral presentation.)
R 161
141
Antlers may regenerate from persistent neural crestlike stem cells.
J. G. Mount, M. Muzylak, S. Allen, S. Okushima, T. Althnaian, I. M. McGonnell, and
J. S. Price
The Royal Veterinary College, London, UK.
Complete regeneration of mammalian organs is currently confined to the natural
annual re-growth of deer antlers. These are shed each spring then re-grow by
endochondral ossification from an undifferentiated antler bud, which forms on the
pedicle (a permanent extension of the frontal bone), into large branched bones. There
is histological evidence that the pedicle periosteum is potentially a source of stem cells
that contribute to formation of the regenerating antler. In this study we explore the
hypothesis that these cells are a population of persistent neural crest-derived stem
cells. Neural crest cells (NCCs) were traditionally thought to be a transient cell type,
present only in embryos. However, multi-potent cells showing NCC characteristics
have recently been identified in adult rodent peripheral tissues. Here we show that
during the antlers regenerative phase, cells in the pedicle periosteum and in the antler
bud display a number of characteristics of NCCs and are multipotential. Using RT-
PCR we detected the expression of a number of NCC markers including slug.
Quantitative RT-PCR showed that slug expression was significantly higher in
periosteum during the regenerative phase compared to the non-regenerative phase.
Immunocytochemistry confirmed that slug localised in progenitors in the antler bud
and in the periosteum of regenerating antlers. Slug also localised in skin during antler
bud re-epithelialisation. However, slug protein was barely detectable in pedicle tissues
during the non-regenerative phase. Multipotentially was demonstrated by culturing
pedicle periosteal cells under appropriate conditions and could be induced to
differentiate into osteoblasts, chondrocytes and adipocytes. These observations
suggest that the regenerating antler is supplied with cells that show characteristics of
embryonic NCCs. These cells appear to persist in a niche in the pedicle periosteum
left by shedding the previous antler. That these cells should be implicated in the only
example of organ regeneration in mammals suggests that, if provided with appropriate
environmental cues, such cells in species other than deer could similarly be induced
to regenerate other organs.
162 R
142
Stem cells isolated from the regenerating antler express key
markers of the osteogenic lineage.
J. Napp
1
, K. G. Wiese
1
, U. Kierdorf
2
, H. Kierdorf
2
, N. Seymour
1
, H. Schliephake
1
, and
H. J. Rolf
1
1
University Hospital, Department of Oral and Maxillofacial Surgery, Clinical
Research, University of Goettingen, D-37075 Goettingen, Germany
2
Department of Biology, University of Hildesheim, D-31141 Hildesheim, Germany
For years, an increasing number of in vitro - studies have dealt with the influence of
hormones and growth factors on the growth of cells derived from the proliferation
zone of regenerating antlers. Presently the role of androgens in the in vitro
proliferation of antlerogenic cells is still controversial. We investigated cell cultures
containing a mixed cell population derived from selected layers of red deer (Cervus
elaphus) and fallow deer (Dama dama) antler tissue. Cultures were treated with
testosterone (T) and dihydrotestosterone (DHT) to investigate, whether different doses
of steroids can influence antler cells that were incubated in serum-containing medium.
We used different cell passages derived from fresh and short-term frozened antler
tissue. In addition, long-term cultures of fallow deer antler cells were also tested. Our
experiments revealed that in both species ''physiological'' doses of testosterone and
DHT (0.5 to 10 ng/ml) exert mitogenic effects on mixed populations of antler cells in
the presence of 10% FCS. For example, in long-term cultivated fallow deer cultures
we observed significant differences between untreated controls and cultures containing
testosterone concentrations of 0.5 ng/ml and 1 ng/ml. With regard to DHT
concentrations we detected the minimum threshold concentration leading to DHT
effects on fallow deer antlerogenic cells in vitro is between 0.5 and 1 ng/ml. Our data
indicate that the observed mitogenic effects of T and DHT might be altered by one or
even more as yet unknown serum factor(s). From our point of view, the observed
mitogenic effect of androgen treatment seems to depend on the stage of differentiation
and organisation of the antler cells, and might be dependent on serum-containing
culture medium.
This study was supported in part by the DFG (grant no RO 2520/1-1).
(Oral presentation.)
R 163
143
Mitogenic effects of androgens on mixed antler cell cultures.
H. J. Rolf
1
, K. G. Wiese
1
, G. A. Bubenik
2
, L. Bartoลก
3
, R. Kotrba
3
, I. Lรผtjens
1
, and H.
Schliephake
1
1
University Hospital, Department of Oral and Maxillofacial Surgery, Clinical
Research, University of Goettingen, D37075 Goettingen, Germany
2
Department of Integrative Biology, University of Guelph, Guelph, Ontario. Canada
N1G 2W1
3
Ethology Group, Research Institute of Animal Production, Praha 10 - Uhลรญnฤves,
Czech Republic
For years, an increasing number of in vitro - studies have dealt with the influence of
hormones and growth factors on the growth of cells derived from the proliferation
zone of regenerating antlers. Presently the role of androgens in the in vitro
proliferation of antlerogenic cells is still controversial. We investigated cell cultures
containing a mixed cell population derived from selected layers of red deer (Cervus
elaphus) and fallow deer (Dama dama) antler tissue. Cultures were treated with
testosterone (T) and dihydrotestosterone (DHT) to investigate, whether different doses
of steroids can influence antler cells that were incubated in serum-containing medium.
We used different cell passages derived from fresh and short-term frozened antler
tissue. In addition, long-term cultures of fallow deer antler cells were also tested. Our
experiments revealed that in both species ''physiological'' doses of testosterone and
DHT (0.5 to 10 ng/ml) exert mitogenic effects on mixed populations of antler cells in
the presence of 10% FCS. For example, in long-term cultivated fallow deer cultures
we observed significant differences between untreated controls and cultures containing
testosterone concentrations of 0.5 ng/ml and 1 ng/ml. With regard to DHT
concentrations we detected the minimum threshold concentration leading to DHT
effects on fallow deer antlerogenic cells in vitro is between 0.5 and 1 ng/ml. Our data
indicate that the observed mitogenic effects of T and DHT might be altered by one or
even more as yet unknown serum factor(s). From our point of view, the observed
mitogenic effect of androgen treatment seems to depend on the stage of differentiation
and organisation of the antler cells, and might be dependent on serum-containing
culture medium.
This study was supported in part by the DFG (grant no RO 2520/1-1).
(Oral presentation.)
164 R
144
Antler growth in red deer stags (Cervus elaphus) depends on
testosterone, but not IGF-1, LH, prolactin or cortisol.
L. Bartoลก
1
, D. Schams
2
, J. ล iler
1
, S. Losos
1
, and G. A. Bubenik
3
1
Ethology Group, Research Institute of Animal Production, Praha 10-Uhลรญnฤves,
Czech Republic,
2
Institute of Physiology, University of Munich-Weihenstephan, Freising-
Weihenstephan, Germany,
3
University of Guelph, Guelph, Ontario, Canada
The role of androgens and IGF-1 in antler growth has been a matter of controversy.
Therefore we tested which hormone is associated with antler growth. Twelve red deer
stags (Cervus elaphus) aged 4 to 9 years were blood sampled and the length of their
velvet antlers was measured in one week intervals ranging from the time prior the
antler casting to the end of antler growth and velvet shedding. Concentrations of
testosterone (T), IGF-1, LH, prolactin and cortisol were determined by RIA The
increase in antler length and hormone concentrations were determined for each
sampling period. The statistical analysis, using a generalized linear mixed model
(GLMM) procedure in SAS, was performed only for samples showing detectable
antler growth. We found that antler growth (an increase of antler length per day) was
dependent on changes in T concentration per day (GLMM, F
(1,42)
=15.94, P=0.0003),
the month when samples were taken (GLMM, F
(4,42)
=11.06, P<0.0001) and the
interaction between age of the stag and changes in T concentration per day (GLMM,
F
(3,42)
=9.56, P<0.0001). The increase in T concentrations and the intensity of antler
growth tended to be higher with age. No significant dependency was established
between the rate of antler growth and the changes in concentrations of any other
hormone investigated. This finding further supports the concept that the range of low
(permissive) concentrations of androgens stimulate antler growth. This does not
challenge the fact that higher levels stop the growth and induce antler mineralization.
(Oral presentation.)
145
Fetal differentiation of the antler developing area in red deer (C.
elaphus).
P. M. F. Audenaerde and P. J. M. Simoens
Marquettepolder 1, Lapscheure 8340, Belgium
At the site of prospective antler development, differentiation of the skin and
subcuteneous tissues was observed both macroscopically and histologically in unborn
R 165
red deer calves. Important characteristics of this area include an increased
pigmentation and epidermal folding in fetuses of both sexes, and local swelling caused
by an accumulation of oedematous mesenchymal subcutaneous tissue which was
obvious in most male fetuses. The different cell types involved in these features were
studied histologically. The localisation of apoptosis was evaluated by means of in situ
end labelling (TUNEL) and androgen receptor distribution was demonstrated by
means of immunohistochemical methods.
(Oral presentation.)
146
Central vessels in roe deer antlers (Capreolus capreolus) - a
histomorphological study.
H. J. Rolf
1
and C. H. Lohmann
2
1
University Hospital, Department of Oral and Maxillofacial Surgery, Clinical
Research, University of Goettingen, D-37075 Goettingen, Germany
2
University Hospital Eppendorf, Dept. for Orthopedics, D-20246 Hamburg, Germany
For decades, the polished deer antlers have been considered ''dead bone''. However,
recently it has been demonstrated that the spongy core and the compact bone of hard
fallow deer (Cervus dama) antlers exhibit a well-developed system of capillaries and
blood vessels connected to the body vascular system via the pedicle. Furthermore,
histological analysis and physiological experiments with fluorescent markers revealed
that main parts of the fallow deer antler bone are still alive at least three weeks prior
to antler casting (Rolf and Enderle, 1999). In order to test our hypothesis that a
vascular blood supply of polished antlers is a general principle in deer we examined
10 roebucks at the age of 1 to 8 years. Eight skulls had polished antlers and two were
complete heads with velvet antlers. We investigated the manner in which the antlers
could be internally supplied with blood vessels. Our specimens revealed definitive
evidence that roe deer antlers also have a similar developed system of capillaries and
vessels as observed in fallow deer. In addition, in roe deer we detected canals passing
internally from the frontal bone pedicle to the antler base and up to the upper parts of
the antler. Besides we have hints that roe deer also possess vessels penetrating the
pedicle bone below the coronet and then passing intramedullary to supply the polished
antler. There were no differences in the presence of blood vessels varying with age.
Thus, we suggest that the detected central vascular canals are main part of a vascular
system that enables blood supply of the antler bone in roe deer even after velvet
shedding. Based on our results it might be possible that the morphology of these
vascular systems supplying polished deer antlers could reflect an interspecies
difference in deer. Whether the central vascular canals in roe deer contain either
arteries or veins remains to be elucidated. Further studies to clarify these questions are
in progress. However, these observations do not seem to be age-dependent in
roebucks.
166 R
References: Rolf, H. J. and Enderle, A. (1999) Hard fallow deer antler: A living bone
till antler casting? Anatomical Record 255:69-77.
(Oral presentation.)
147
Antler characteristics of the Sardinian red deer (Cervus elaphus
corsicanus): a preliminary analysis.
A. Caboni, C. Murgia, and S. Mattioli
Department of Animal Biology and Ecology, University of Cagliari, Viale Poetto,
Viale Poetto 1, 09100 Cagliari, Italy
Since the first descriptions were published in the XVIII
th
century, Sardinian red deer
has been known for its small body size and simple antler design. Here for the first time
the main morphometric traits of antlers from the three subpopulation surviving in
southern Sardinia were analysed. We have examined a total of 216 antler beams
belonging to stags 2 years and older: 49 antler pairs are from animals found dead and
118 are cast antler beams. Considering adult stags (5 years old), the most frequent
structure is the four-tined beam (brow tine, trez tine and terminal fork), with an
incidence of 52.2 % (n= 157). Simple spikes, i.e. unbranched antler beams, account
for 0.6% of total beams. The maximum number of tines per beam is invariably 6.
Twelve-tined antler pairs represent only 2.1%. Bez tine is rare, with a frequency of
just 7.3%. Crown is uncommon, occurring in 14 % of beams and far from the typical
cup-like shape. A palmation of the upper parts of the antler is a relatively frequent
trait, occurring in 18.7 % of adults. The longest antler beam is 85 cm, the heaviest 1.2
kg, the averages being about 63 cm and 0.6 kg.
In subadults (2-4 years old) antler beams with three tines prevail, with an incidence
of 36.2% (n= 58). Spikes account for 29.3% of total beams. Palmation occurs in
5.2 %, while bez tine and crown are always absent. The antler conformation and size
of Sardinian red deer documented by our survey fit well within the maintenance
phenotype model described by Geist, substantially comparable to those of red deer
from North Africa, southern Spain, Scottish Highlands living in scarcely productive
habitats.
(Oral presentation.)
R 167
148
What we can learn from antler composition and structure: from
nutrition to management.
T. Landete-Castillejos, J. A. Estรฉvez, A. J. Garcia, F. Ceacero, E. Gaspar-Lรณpez, D.
Carriรณn, and L. Gallego
IREC, University of Castilla-La Mancha, Albacete, Spain
Researchers have devoted little attention to bone chemical composition and this has
been considered basically constant under normal nutrition conditions. However, if
antler bone composition proved to show variation associated to antler characteristics,
body size or physiological state, and recent diet, assessing composition and structure
might be a useful tool for population diagnosis and management. This talk will discuss
a recent line of studies showing factors affecting antler bone composition and internal
structure. In Iberian red deer spikes, content of 25 antlers was assessed for ash, Ca, P,
K, Na, Mg, Fe and Zn in base and tine to explain variability in antler length, weight,
and perimeter. In turn, mean composition and difference in concentration of each
mineral was related to body measures at one year of age, weight at birth, that at one
year of age and gains during lactation, or between weaning and year of age. Chemical
composition differed between base and tine in ash, Ca, P, K, Zn and Fe, but not in Na
or Mg. Composition explained a range of variability from 76 to 99% in antler
characteristics. Body weight and size, in turn, influenced mineral composition. The
greatest body effect was that of gains during lactation on most macro-minerals, which
explained a mean variability of 50%. Another study showed that antler bone
composition in a captive population with high plane of nutrition and parasite/pathogen
treatments differed from those from a free-ranging population with lower quality food
and no parasite/pathogen treatments. In addition, cortical layer of antler in antlers of
deer living in controlled conditions was thicker than in those of free-ranging deer.
Finally, mineral content and opacity to X-rays was also greater in the former than in
antlers of free-ranging deer. In conclusion, antler bone composition is variable in
normal conditions, variability appears to depend on region of the antler and
maintenance conditions, and this together with structural differences suggest that both
might play a role in biomechanical properties of the antler. It might also show the
nutritional status or physiological effort to grow antlers.
(Oral presentation.)
168 R
149
Post-velvet shedding antler histology of red deer (Cervus elaphus)
living in the wild.
A. Dobrowolska and K. Gรณrecka
Department of Animal Anatomy, Szczecin,Poland
When red deer stags fight, parts of their antler beams may break off. Two right beams,
broken off beneath the tray tine, were found in late September, just after stag fights
terminated. One of the beams terminated with crown tines, the other being forked.
Both beams bled at the site of breaking and exuded a strong, specific smell,
perceivable for a few days. Most probably, the smell was related to the heat period and
the resultant high content of sex hormones in stag blood. This is evidence that blood
circulation in the antler cortical part is maintained, even after the velvet has been shed.
The beam histological structure was analysed. The aim of the study was to provide
observations on the structure and blood vessel system of the antlers, after the velvet
has been shed, in the red deer living in the wild, a few months prior to physiological
antler casting. Histological slides were made in the form of tranverse sections of the
beams to provide materials for observations on antler structure in the cortical and core
part. The beams were cut at various heights to yield 20 transverse sections each. When
viewed under the microscope, the antler cross sections were observed to contain bone
lamellae, concentrically arranged around Haversian canals, which formed osteons of
various diameters; there were also interstitial lamellae, located inbetween the
neighbouring osteons. The osteal and interstitial lamellae were interspersed by bone
cavities containing osteocytes. The cortical part of the antlers was formed by densely
packed, fairly regular, circular and oval osteons; their diameters weresmaller than
those in the core part, their Haversian canals being smaller as well. On the other hand,
the antler core was formed by osteons consisting of a lower number of osteal lamellae
and large, frequently irregular Haversian canals which showed fragments of blood
vessel with blood remains (some canals were entirely filled with blood). The
Haversian canals, both in the cortical part and in the core, gave rise to numerous
Volkmannโs canals. Some Haversian canals, particularly in the antler core, showed a
clear polarity. The pole of such Haversian canals was found to contain active
osteoblasts. The remodelling of bone tissue was particularly well visible in the core,
the remodelling continuing after velvet shedding and involving the formation of new,
small osteons at the ends of Volkmannโs canals, at the junction of at least three
osteons, between the interstital lamellae.
(Poster presentation.)
R 169
150
Lengths of pedicles and antlers in Reeves' muntjac.
N. G. Chapman
Larkmead, 29 The Street, Barton Mills, Bury St.Edmunds, UK
Most species of deer have long antlers on short pedicles which arise from the frontal
bones. The pedicles do not reach full size until 14-16 months of age. In contrast,
muntjac have long pedicles, arising from a ridge at the supraorbital margins, and short
antlers. This is the major reason for placing muntjac in a separate family, Muntiacidae.
In all deer the dimensions of the pedicles can change over the years as remodelling of
bone takes place. Using a large sample of skulls of Reeves' muntjac(Muntiacus
reevesi), some of exactly known age, the lengths of the pedicles and antlers of first
and subsequent heads will be compared.
(Poster presentation.)
151
Consistent interindividual variability in proliferation potential of
antler cells cultivated in vitro under various treatments.
E. Kuลพmovรก
1, 2
, L. Bartoลก
1
, M. Tomรกnek
1
, R. Kotrba
1
, and G. A. Bubenik
3
1
Research Institute of Animal Production, Prague 10-Uhลรญnฤves, Czech Republic
2
Department of Ecology, Charles University, Prague, Czech Republic
3
Department of Integrative Biology, University of Guelph, Guelph, Ontario, N1G
2W1, Canada
Deer antlers and their growth are under the control of many external and internal
factors. These mechanisms are still not sufficiently clarified neither at physiological,
cellular, nor molecular level. Antlers periodically regenerate and their growth is one
of the fastest in the animal kingdom. The antler cells have a great proliferation
potential. It has been postulated over the last two decades that the antler growth and
the proliferation of antler cells is influenced by steroid hormones and/or growth
factors such as the insulin-like growth factor I (IGF-I).
In this study, antler cells were derived from the tips of regenerating antlers of three-
year old red deer stags (A, B, C). Primary cultures and first passages were exposed to
various experimental procedures with 1% and 10% foetal bovine serum (FBS) and a
mixture of 1% FBS supplemented with oestradiol, testosterone or IGF-I. The
proliferation potential of these cells was determined by incorporation of tritium-
labelled thymidine. Consistent significant differences have been reported among
individual animals across all experimental procedures using the GLMM model (PROC
MIXED in SAS). Stag A exhibited the highest while stag B the lowest values in all
experiments. Values of the stag C were between those of stags A and B, usually closer
170 R
to B rather than to A. The average values of A were 2.1 to 20.1 times higher than
those of individual B (P<0.01 to P<0.0001). The results suggest that the individual
potential of each animal may significantly affect the cultivation of antler cells and
hence should be reported in all in vitro experiments.
(Poster presentation.)
Responses of deer to global
environmental change
172 R
152
Biogeography of Cervidae in Peru.
J. Barrio
University of Florida and Universidad Nacional Agraria La Molina, Luis Garcia
Rojas 175, Urb. Humboldt, Miraflores, Lima 18, Peru
Peru is one of the most biological diverse countries in the world thanks to its
geography and its variety of climates. When considering deer species, Peru contains
both tropical and temperate forms, including five of the six genera native to South
America. Of the 13 recognized native species from South America, seven were known
to occur in Peru (Blastocerus dichotomus, Hippocamelus antisensis, Mazama
americana, M. chunyi, M. gouazoubira, Odocoileus virginianus and Pudu
mephistophiles). However, new data and collected material confirmed the presence of
Mazama rufina, beforehand mistakenly included as present due to confusion with a
very small subspecies of M. americana. Also, an undescribed species has been
photographed in the Amazon lowlands, and probably the small M. americana
subspecies may warrant specific status given large morphological differences. Here,
data on distribution and habitats used are presented for all species occurring in Peru.
The knowledge on the distributional ranges of Mazama rufina and Pudu mephistophile
are here improved, in the first species increasing its range southwards, and in Pudu
closing the gap between the northern population, formerly known only from Colombia
and Ecuador, and the southern Peruvian population. Also, habitat use differences
between Hippocamelus antisensis and Odocoileus virginianus peruvianus, and
between Mazama americana and Mazama gouazoubira nemorivaga are described.
Hippocamelus and Odocoileus seemed to separate themselves by altitude and habitat
type, whether Mazama species presented a complex division where areas dominated
by specific habitat types showed notorious higher densities of either species over the
other. M. gouazoubira nemorivaga , were more abundant in areas with the canopy
dominated by Brazil nut (Bertholetia excelsa), and areas where the undergrowth was
dominated by the short palm Irapai sp. M. americana was the only species found in
seasonally inundated lowland forest. South American deer species present complex
niche divisions between species and distributional overlap among several species. The
overlap among species in Peru is most notorious throughout the montane and cloud
forests. Therefore, there are several macro habitat and biogeography concerns still
unexplored that are starting to be understood, that need studies before being
encroached by and lost to the growing agricultural frontier.
(Oral presentation.)
R 173
153
The influence of season, food intake, and social rank on cortisol
secretion in red deer (Cervus elaphus).
F. Balfanz
1
, C. Beiglbรถck
1
, S. Huber
1
, R. Palme
2
, and W. Arnold
1
1
Research Institute of Wildlife Ecology, University of Veterinary Medicine, Vienna,
Austria
2
Institute of Biochemistry, University of Veterinary Medicine Vienna, Austria
Deer developed as an adaptation to the nutritional bottle-neck of winter pronounced
cycles of voluntary food intake, body mass, and metabolic rate. We investigated in red
deer how this seasonal adaptation is reflected in the secretion of glucocorticosteroids
measured as the concentration of glucocorticoid metabolites (GCM) in faeces, and
how food availability influences the annual rhythm. We sampled faeces throughout
the year from a captive population living under close to natural conditions in a large
study enclosure, and from free-living deer. The captive deer had during one study year
during winter access to a limited amount of hay and beets in addition to natural forage,
during two other study years to pellets ad lib. throughout the year. In addition, we
studied the effect of social interactions on GCM secretion. GCM secretion varied
periodically with a peak during winter and a nadir during summer. The amplitude of
this seasonal cycle was on average about 10% of the annual mean both in the captive
population receiving only limited supplemental food during winter and in the natural
population. Under ad lib. feeding the amplitude of mean GMB secretion was
attenuated to 5% of the annual mean. GMB secretion was further influenced by social
rank with highest values found in individuals of intermediate rank. We conclude that
the amount of glucocorticosteroids secreted is an important endocrine signal in the
regulation of the annual change from an anabolic growth phase timed to summer when
food is abundant with an accumulation of body energy reserves, to a winter
physiology characterized by a catabolic metabolism and anorexia. This cycle is
controlled by an endogenous rhythm entrained to the natural photoperiod that enables
anticipation of the seasonal environmental changes and timely adaptation. However,
the magnitude of the reaction and presumably to some degree also its phase is
modulated by food availability. Our results further demonstrate an important influence
of social rank on GMB secretion. However, glucocorticoid and GMB secretion can
be used as reliable indicators of stress only when controlling for the confounding
effect of season and food availability. For assessing the latter, variation of climate,
habitat quality, and population density needs to be considered.
(Oral presentation.)
174 R
154
Defense of territories by rutting red deer stags, Cervus elaphus, in
Patagonia, Argentina.
J. M. Smith-Flueck and W. T. Flueck
Arelauquen Foundation C.C. 176, 8400 San Carlos de Bariloche, Argentina
Introduction
Mating behavior of both Old and New World Cervus elaphus males has generally
been characterized as harem defense (Struhsaker 1967, Clutton-Brock et al., 1982),
basically referring to mobile harems where females are followed and defended across
their foraging range by rutting males. An alternative strategy whereby males establish
mating territories has been previously described for red deer populations in the
savannah-like Pampas of Argentina (Dietrich 1987) and in southern Iberia (Carranza
et al., 1990). Defense of mobile harem and mating territories by prime males,
evidencing intraspecific variation in red deer mating systems, are only known to
coexist in southern Iberia (Carranza et al., 1990, 1995). Our study, a first to
empirically describe mating behaviors of territorial red deer in Argentina, was
conducted to determine if the mating strategies employed by prime stags of the
Andean-Patagonian cordillera were comparable to that of the southern Iberian
population, as both populations enter the rut at the end of the dry season, when forage
availability is at its lowest and more patchily distributed across the landscape. If
resource distribution indirectly influences choice of mating systems (Carranza 1995),
one might also expect to find rutting stags holding territories in the lower elevations
of the ecotone habitat of the Andean cordillera, where high numbers of females
concentrate in open areas to forage during the rut. This study tested if dominant rutting
males in areas with high female concentrations were defending territories. The various
mating strategies employed by rutting males of this Andean cordillera population are
described, providing a new example of between-population variation of mating
systems.
Study Area
The study population inhabits the mountainous ecotone and steppe habitats of the
eastern Andean cordillera in Patagonia where the dominant climate is temperate with
main precipitation occurring between April and September. The breeding season,
occurring sometime between early March and late April, lasts 3 to 6 weeks, and peaks
around the last 2 weeks of March. The study area was located on private land within
the Nahuel Huapi national reserve, (40158"S; 71112"W), Argentina. The area over
which the rut takes place is characterized by a mosaic of habitats:
1) forest patches represented by a variety of pure and mixed stands of Nothofagus
antarctica, Austrocedrus chilensis, Lomatia hirsuta, Maytenus boaria and Schinus
patagonicus;
2) brush patches represented predominately by Berberis spp., and Colletia
spinosissima.
R 175
3) mallins - a type of wet grassland - common throughout the eastern cordillera of the
Andes, which are defined as exhibiting running water or inundated soils all year
round. Cyperaceae, Juncaceae and other wetland species dominate the wet central
sections of the mallins. Of the graminoides, Carex canescens and Carex macloviana
(Cyperaceae) and Juncus depauperatus are the most prevalent species. The dry areas
of the mallin have more xeric graminoid and forb species, such as Festuca pallescens,
Hordeum chilense, Poa spp, and Ranunculus peduncularis;
4) grass-dominated steppe Stipa speciosa var. major and Festuca pallescens with
variable occurrence of brush species like Mulinum spinosum, Berberis spp., and
Colletia spinosissima;
5) riparian habitat with Salix sp. and Nothofagus antarctica the predominate species.
General observations on the rut behaviour were made over the entire range of habitat
types, however, the recorded observations to test male breeding behavior were
conducted from a blind overlooking a mallin (total area of 5 ha). Patches of shrubs and
a few singular trees dotted the peripheral areas of the mallin, and consisted mainly of
Berberis spp., introduced European Rosa mosquesta, Nothofagus antarctica, Lomatia
hirsuta, Maytenus boaria, and a few ancient apple trees, Malus pumila.
Methods
Binoculars and scopes were used to observe red deer rutting behaviour from
sunrise to sunset from March 9 through April 9, 1995, using ad-libitum sampling and
focal-animal sampling techniques Altmann (1973). The focal sampling was based on
a minimum recording period of 20 minutes. Additionally the study area was scanned
once each hour to record total number of individuals present and the age and sex
category of each. Number of males holding territories were noted along with harem
size and location. Location of males throughout the observation period were recorded
on a map with grids to determine temporal use of the area. Sketches of antler
configurations for each territorial male helped to distinguish individual males on the
study site, later verified with photographs. Vocalization was also used to distinguish
locations of males using the mixed brush/forest patch habitat adjoining the mallin.
Length of stay, location on study site and dominant status of each identified male on
the mallin was determined whenever data allowed.
Males were randomly selected on the mallin to conduct focal-animal scans
between the hours of 800 to1100 and 1430 to 2030, corresponding to the daylight
hours with highest deer activity. Behavior recorded during these scans were divided
into agonistic male-to-male interactions (displacing, chase, thrash, parallel walk,
initiating and sparring) and male-to-female herding and courting behaviour (sniffing,
licking, flehman, chivying, and mounting) as described by Clutton-Brock et al., (1992)
and Geist (2002). Although all occurrences of these behaviours were recorded during
the entire focal-animal sampling period, for the statistical tests performed, more than
one observation of a particular behaviour performed by the focal animal during a 1-
minute unit of observation was tabulated as a single occurrence.
Additional observations were made of daily movement patterns of hinds between
the mallin and brushy/forested slopes to the east and west of the main study site.
Males responding to the females were noted along with a description of their call, and
body and antler size.
To distinguish whether males were defending territories or females, each focal
176 R
sampling of an individual male, being the mean of several 1-minute units of
observation (Carranza et al., 1990), was used to conduct a Student t-test to compare
differences in interactions between sedentary males with and without females. To
evaluate if the intensity of rut behaviour of focal males might be affected by the
number of females in their harem, a Spearman rank correlation was performed on
focal sampling data to compare number of females in harem with mean frequency of
male-to-male interactions, and also to compare number of females in harem with mean
frequency of male-to-female interactions.
Results and Discussion
The population density for the study year, estimated from pellet count transects,
was approximately 100 deer per km
2
. Males were found to defend territories ranging
from 0.8 to 2.5 ha in size. The territorial boundaries established by the focal males
shifted occasionally between successive hours and days, similar to observations of
Iberian red deer males (Carranza et al., 1990). Number of territorial males in mallin
at any one time varied from 0 to 6. Only males of larger body and antler mass held
territories (n=13). Moreover, throughout the peak period of the rut, our focal males
would remain the entire day in the same general area of the mallin, leaving only if
disturbed. Males remained in established territories, even when females were absent,
often bedding down in the heat of the day, when calling would continue, though
sporadically and with less intensity. An occasional intruding male was cause for the
bedded males to rouse, concomitantly increasing the frequency of their calls. The
roaring activity also picked up in the late afternoon, stimulated by females entering the
mallin. Timing of activity patterns varied daily according to the meteorological
conditions.
During the peak of the rutting period, a male=s breeding status was influenced by
his body mass and antler size (Clutton-Brock et al., 1982). The larger dominant males
remained on the mallin through the peak of the rutting period, while the smaller
subordinate males remained at the transition zone bordering the mallin. Occasionally
these satellite males entered the mallin, causing the dominant territory holders to
engage in defense activity. The youngest age class males were the most mobile,
moving in and around the territories of the older males. An individual male=s strategy
sometimes changed during the course of a breeding season due to a social status
change or when a territorial site suddenly becoming vacant. If a dominant male, for
instance, left the main breeding area to migrate to his winter range, a less dominant
stag would replace him. Often males beyond the prime breeding age would move into
these sites. It is conceivable that these males, during their prime years, may have held
territories in this same mallin.
Males of subordinate status, unable to compete with the bigger males on the
mallin, adopted an alternate strategy by taking advantaging of the femaleโs daily
movement patterns. Although some females might remain on the open mallin
throughout the entire day, many females returned to the brushy forested slopes of the
hills to the east and northwest of the main study site to bed down. Once feeding
activity commenced in the late afternoon, the position of females groups moving
slowly down the vegetated hillside was made evident by males calling as the females
approached and passed by their location. These males would follow the female group
a short distance, as far as to the next waiting male. Roaring activity was thus highest
R 177
on the hillside during the ascent and descent of females in the morning and evening
hours, respectively. These subordinate males remained loosely in the same general
area along the hillside, with highest densities of males along the lower portion of the
slopes near the transition to the mallin. Here was the waiting zone, where satellite
males stood ready for transient females to pass on their way to the mallin. This mating
strategy would benefit smaller males, allowing them to avoid conflicts with males of
larger stature and yet provide them with opportunities to interact with potential estrous
females. At the beginning and tail end of the rut, when densities of males were lower
on the open mallin, movement by these less dominant stags was greater.
Rates of male agonistic behaviour during the focal-animal samplings varied
greatly, triggered by various factors such as occasional male intruders or females
crossing the territory. Although frequency of a male=s aggressive interactions towards
other males was not affected by the number of females in his harem (r
S
= -0.15, n =
37, NS), his frequency of interactions with females positively correlated with number
of females in the harem (r
S
= 0.50, n = 37, p = 0.002). A focal sedentary male reacted
aggressively to other males regardless of the presence of females within his territory
(mean " SD number of defensive activities per min: 0.058 " 0.075 with females
versus 0.096 " 0.145 without females; t = 1.108, df = 42, NS). Such defensive
behavior by the males in the absence of females, which concurs with results of
Carranza et al., (1990), indicates that males were not defending a harem but rather
their area.
Resource distribution has been suggested as a factor influencing choice of mating
systems, afforded by behavioral plasticity (Carranza 1995). In northern Europe, where
the main mating strategy for stags is the defense of mobile harems (Clutton-Brock et
al., 1982), general food availability is still high during the rut so that females are not
forced to clump at particular feeding sites. In contrast in southern Iberia, the mating
season takes place during the period of lowest food availability, after the dry and hot
summer. Whether the underlying mechanisms for the unique behavioral repertoire
observed for the red deer stags in the eastern Andean cordillera are a function of
habitat structure (environmental heterogeneity and patchy resource distribution),
population densities, herd composition, group size, genetic makeup or combinations
of these factors remains to be determined. Studies comparing intraspecific variation
of mating strategies between and within populations can lend insight into the
functioning mechanisms. It is conceivable that strategies employed by rutting stags
during the study period, when the density of the study population in the ecotone was
about 100 deer/km
2
, would alternate once densities are driven down as was shown for
lekking fallow deer (Apollonio 1989). The suggestion by Carranza et al., (1995) that
territoriality should arise where resources are scarce and patchy during the rut is
supported by findings of this study, in that ecosystem resources, though not scarce like
in Iberia, are patchily distributed. The heterogeneous landscape, affecting deer
distribution, allows females to use various habitat types during the rut, with large
group concentrations in the open mallins and grasslands. Their movement patterns
appeared to influence the strategy selected by the males, which was in turn affected
by a male=s dominance status, a reflection of his body and antler mass. This study, a
first to describe mating behaviors of territorial male red deer in Patagonia, did not find
any evidence of mobile harem defense in the habitat types studied; instead, territorial
178 R
defense, in open areas receiving high female concentrations, was the strategy of choice
by the prime breeding males. Non-prime males exhibited various other semblances of
mating strategies. Casual observations of rutting by prime males occurring in other
habitat types, like closed forest, indicate that several strategies are being employed
across this heterogenous landscape. Further investigation focusing on more
homogeneous parts of the range, such as the Andean montane grasslands or grass-
dominated steppe habitat might reveal males utilizing mobile harem defense. The
repertoire of mating strategies employed by prime red deer stags in this Patagonian
population appears to show that intraspecific variation in deer mating systems may be
more common than previously recognized.
References
Altmann, J. (1973) Observational Study of Behavior Sampling Methods. Behavior 49:
227-267.
Apollonio, M. (1989) Lekking in fallow deer: just a matter of density? Ethology
Ecology & Evolution 1: 291-294.
Clutton-Brock, T.H., Guinness, F.E., Albon, S.D. (1982) Red deer: behavior and
ecology of two sexes. The University of Chicago Press, Chicago.
Carranza, J., Alvarez, F., Redondo, T. (1990) Territoriality as a mating strategy in red
deer. Animal Behavior 40: 79-88.
Carranza, J., Garcia-Munoz, A.J., Dios Vargas, J. de (1995) Experimental shifting
from harem defense to territoriality in rutting red deer. Animal Behavior 49:551-554.
Geist V. (2002) Adaptive behavioral Strategies. Pages 389-447 in D.E. Toweill and
J Ward Thomas, editors. North American elk: ecology and management. Smithsonian
Institution Press, Washington.
Struhsaker, T.T. (1967) Behavior of elk (Cervus canadensis) during the rut. Zeitschrift
fรผr Tierpsychol. 24: 80-114.
(Oral presentation.)
155
Spatial behavior paths of food search in roe deer (Capreolus
capreolus).
S. Said, M. Pellerin, M. Le Corre, O. Widmer, and G. Van Laere
ONCFS-CNERA CS, 55000 Bar Le Duc, France
The way roe deer exploit resources within their home range is not well known because
of cryptic behaviour and dense habitat. Furthermore, radio tracking locations used to
R 179
estimate home ranges are insufficient to obtain animal paths. However, the appearance
of GPS (Global Positioning System) collars permits a greater number of positions and
therefore the determination of movement patterns. Several methods have been
developed to characterize habitat structure and animal movements. One of these, the
First Passage Time (Fauchald and Tveraa 2003), can be used as a measurement of
search effort along a path and is defined as the time required for an animal to cross a
circle with a given radius. We applied this analysis to GPS locations of roe deer
females in Chizรฉ and Trois-Fontaines Forests (France) to understand how animals
change their movement patterns in relation to the environment (autumn 2003: N =
2+5, and spring-summer 2004: N = 9+9). Our results revealed circles of intensive
searching measuring 10 to 100 meters, and for every female the majority of zones
appeared in all paths. Furthermore, preliminary results about animal activity in these
circles, provided by GPS collars, suggested that for some females these were rest
zones rather than zones of high activity. However, further analyses are necessary and
in progress. Intersections of detected zones with vegetation, i.e. plant species and
biomass and percent of habitat opening, are still in analysis. These additional results
will be presented during the meeting.
(Oral presentation.)
156
Carbon and nitrogen efficiencies in venison production.
M. H. Davies, D. G. Chapple, and B. Cottrill
ADAS UK Ltd, Rosemaund, Preston Wynne, Hereford, HR1 3PG, UK
The UK farmed deer industry is relatively small with some 250 farmers and 33,000
deer. A desk study was undertaken during 2003 to assess the environmental impact
of venison production in relation to the utilisation of nitrogen and carbon. Agriculture
is the main source of ammonia (NH
3
) emission (82% of total emissions), and
contributes significantly to methane (CH
4
) and nitrous oxide (N
2
O) pollution, at 35%
and 64% of totals emissions respectively. Deer are ruminants and are opportunistic or
adaptable feeders with an evolutionary tendency towards fibre digestion (grass and
roughage eaters). They are basically diurnal, usually having 5-11 feeding periods in
a 24-hour period. Deer have a 30% higher fasting metabolic rate than sheep, but
similar to cattle. They have a 38% higher energy requirement for maintenance than
sheep, but the higher heat loss in deer is balanced by more fat deposited in sheep. Deer
exhibit changes in digestion strategies related to season and metabolism, and may
utilise forage with a high content of soluble carbohydrate and protein more efficiently
than sheep or cattle at certain times of the year. Methane production is an inescapable
consequence of fibre digestion by herbivores. The most effective mitigation strategies
to reduce methane emission are likely to be those associated with more intensive
production systems, with reduced feed input per unit of product output. The review
concluded that there is a paucity of data on the emission of ammonia and nitrous oxide
180 R
by deer. Most estimates are derived by assuming similarities with other domestic
animals, with estimates scaled to account for differences in body size. Deer calves
during their first winter are generally fed conserved silage, and more targeted protein
supplementation may improve N utilisation. However, most of the ammonia produced
by deer will be at grass, where there is limited scope for altering the protein content
of the diet, although tanniferous plants may offer some potential to improve N
utilisation. The environmental impact of more deer farming in the UK is difficult to
ascertain. In general, deer are somewhat less efficient biologically in converting feed
to meat, but the niche product they produce, which commands a premium price, does
compensate for this. Our current understanding of the digestive processes in deer does
not indicate it is vastly different from that of other ruminants, and therefore a small-
medium swing to deer farming is unlikely to have a huge environmental impact.
(Oral presentation.)
157
Methane production by farmed red deer.
N. M. Swainson
1
, S. O. Hoskin
1
, and H. Clark
2
1
Institute of Veterinary, Animal and Biomedical Sciences, Massey University,
Palmerston North, New Zealand.
2
AgResearch Ltd, Grasslands Research Centre, Palmerston North, New Zealand
Methane emissions from ruminant livestock in New Zealand are expected to increase
by 10 to 20% between 1990 and 2010, the beginning of the first commitment period
of the Kyoto Protocol. New Zealandโs obligations under the Kyoto Protocol require
the Government to take responsibility for emissions in excess of the 1990 baseline
year. Therefore, methods need to be developed to reduce methane emissions.
However, before such methods can be developed, a better understanding of animal and
nutrition factors affecting methane emissions across ruminant species is needed.
Methane is not only a potent greenhouse gas, but its production represents a loss of
energy from the feed consumed by ruminant animals. Assuming a price of ten cents
per kilogram of pasture dry matter, it is estimated that the energy lost as methane costs
the New Zealand Deer Industry between $5m and $27m per year. Although deer only
produce a small proportion of the total methane emissions from ruminant livestock in
New Zealand, emissions from the deer industry increased by 60.3% between 1990 and
2001. Numerous experiments to measure methane emissions from sheep and dairy
cows have been conducted. However, until recently, there were no direct
measurements of methane emissions from farmed deer in New Zealand. Experimental
work with adult and weaner deer are summarised and these data indicate that methane
emissions from deer are similar to sheep and dairy cattle per kilogram of dry matter
eaten. However, this has not been confirmed by a direct comparison with animals of
different species fed the same diet at the same time of year.
(Oral presentation.)
R 181
158
Why the Patagonian huemul deer in Argentina fails to recover: An
ecological hypothesis.
W. T. Flueck and J. M. Smith-Flueck
CONICET, Bariloche, Argentina and Fundaciรณn Arelauquen, Bariloche, Argentina
Huemul (Hippocamelus bisulcus) had declined drastically by the 20
th
century. First
field studies in Argentina beran in 1988. The 1
st
Chilean-Argentine meeting (1992)
resulted in a preliminary list of hypothetical factors potentially important for huemul
recovery. These factors are still claimed dogmatically to cause their declines and
failure to recover, though without supportive studies. Efforts still aim to reduce
impacts of these factors, but without monitoring, it is not known if these actions are
relevant or successful. Huemul are in a precarious state. Only 350-600 animals remain
along 1800 km of Andes in Argentina, without known cases of numeric responses or
recolonization, yet several populations have vanished in the last few decades, casting
doubt on success of earlier actions [1]. Here we evaluate the importance of these
factors. We posit that major underlying problems explaining the lack of huemul
recovery have not yet been considered: huemul likely are deficient in trace minerals
important for recruitment.
Hypotheses from 1992 are reevaluated. a) Extensive livestock raising: explanations
are needed for populations which coexisted with cattle for >110 years, or which have
declined, are declining or went extinct without apparent contact with livestock. b)
Replacement of native forest with exotic trees: no study shows negative impacts on
huemul. Replacement of native forest by exotics is illegal, and earlier plantations do
not invade native forest. c) Irrational management of native forests: no study shows
negative impacts on huemul. On the contrary, huemul respond positively to areas
formerly logged and burnt, like growing populations in Chile exhibiting the highest
known huemul density. Current successful recolonizations in Chile into treeless areas
corroborate that huemul may be pastoreal and only secondarily adapted to sylvan
habitat. d) Introduction of exotic animals: no study shows negative impacts on
huemul, only that exotics eat the same plants and have similar preferences as huemul
[2]. This is only relevant if it reduces huemul population growth rates. Considering
huemul diet studies, they likely shift diet without necessarily affecting reproduction
[3], as is known for other cervid species. Besides, in the absence of exotics, huemul
went extinct or have declined. Considering high densities of exotics in former huemul
areas, it appears unlikely that such habitat is limiting huemul in terms of energy and
major plant nutrients [3]. Huemul also form multi-species guild assemblages, thus
presence per se of other herbivore species is unlikely to be problematic, as evidenced
by documented coexistence with livestock. e) Illegal hunting: no information exists
about extent or effects on populations. It is likely negligible in remote remaining
182 R
populations, but may prevent migratory behavior. f) Diseases: no study shows
negative impacts on Argentine huemul. Records from local government research
institutions, animal health authorities and veterinarians would indicate the presence
of dangerous diseases near huemul, as livestock raising is of major economic
importance. Of 376 red deer necropsied, the only unusual observation was a yet
unknown exotic Taenia [4]. In Chile, no disease problems have been found in huemul.
Anecdotal accounts by settlers are cited to claim that foot and mouth disease (FMD)
was responsible for decimating huemul over huge areas 60-70 years ago. Based on
behavior of 5 wild cervid species in UK, these are considered unlikely to be important
in maintaining and transmitting FMD. At normal cervid densities, FMD is self-
limiting. Very low huemul densities and reactions of other cervids to FMD makes
those early anecdotal accounts doubtful. Also, considering known growth rates, a
population of huemul in only 6 years would have increased by 300%. Others claim
that Cysticercus tenuicollis, when transmitted by livestock is fatal to huemul, citing
Texera [5]. However, that author did not consider presence of C. t. to be the cause of
death, rather that the condition of the female deteriorated after a premature parturition,
aggravated by very little space and little variety of food provided. Furthermore, in
other cervids and ungulates, presence of C. t. is considered of little significance. g)
Dogs: no study shows negative impacts on huemul. Anecdotal accounts indicate that
huemul is easily killed by dogs, assumed to be related to absence of native cursorial
predators, and that they become paralyzed due to panic, and thus become easy prey.
However, the evolutionary history of huemul included cursorial predators in North and
South America, including Canis. These large canids became extinct in South America
only in the Holocene. Other Odocoilines employ a hide-and-freeze strategy, bolt or
run off at close encounter, and take to water. Mule deer tend to bound uphill, imposing
heavy costs on a predator, whereas white-tailed deer bolt down and along hillsides.
Huemul are known to snort, stomp the ground, they run, trot or race away uphill or
downhill effortlessly; they also bound like mule deer and take to water. Documented
interactions of huemul with dogs are rare, and thus the more instructive (B. Thomas,
pers. com.). Case 1: a huemul buck ran down a logging trail, bounding side to side to
navigate logs while trying to outrun 3 dogs. Case 2: a female with calf were chased
by 2 dogs; they did not run to water but contoured a hill feature for about 1,5 km and
then climbed to the higher ground, outrunning the dogs. Case 3: a radio- collared
huemul doe and her week-old young were observed. The doe suddenly went in front
of several approaching dogs and away from the bedded calf. The barking dogs circled
away from the area to about 500 m. After 20 minutes the barking stopped. About 4
hours later, just at dark the doe approach the fawn from the other direction, nursing
it and then heading back with it the same way she had come, taking the fawn out of
the area some 500 m. Certainly, any loss is important for severely reduced groups,
including from puma predation (Puma concolor), but this is related to small
population size and not to predation per se [6]. h) Small population size, fragmentation
and genetic isolation: although no studies on huemul document population size,
degree of fragmentation (metapopulation dynamics) or genetic isolation, comparative
studies support the importance of these factors when they exist.
What might account for a generalized absence of recolonization or numeric
responses? A growing huemul population in Chile had a lambda of 1,21 and life spans
of at least 14 years [2]. This numeric response occurred in presence of some dog
R 183
predation and natural predators (puma, foxes). As several Chilean populations have
been recovering, but no such cases are known from Argentina, we evaluated the
potential of nutritional factors which likely could affect reproduction and survival
more so on the Argentine than Chilean side of the Andes.
Geology and Pedology: The central Andes occupied by huemul are characterized
by acidic rocks, landscapes strongly modified by past glaciations, and acidic soils
strongly influenced by volcanic depositions, all of which favor low selenium (Se) and
iodine levels. The influence of volcanic ash diminishes to the east due to drier
climates. Westerly Pacific winds result in minimal aerial Se input which is highest by
the coast and decreases further inland. The gradient of iodine depositions also
decreases from west to east, and with increasing altitude in the Andes. Cattle near the
Peruvian coast had levels 6 times higher than found inland at higher altitude. Hence,
Se and iodine provision in Argentina is expected to be lower than in Chile.
Topography modulates soil concentrations as leaching occurs on ridge land and results
in a decrease of soil Se and iodine, while adjacent valleys maintain or increase Se and
iodine levels [7]. Past and current land use further diminish bioavailable Se and iodine
[8].
Biochemical Functions: Only recently discovered, the genetic code had to be
expanded and Se forms part of the 21
st
amino acid, essential in all mammals. Se, an
active part of several enzymes, is involved in oxygen metabolism and thus functions
at very basic biochemical levels, and deficiency is thus expressed in many ways. In
all animal species studied, deficiency impairs reproductive performance of females
and males. In ruminant neonates, white muscle disease is typical of deficiency. It also
is fundamental for proper immune function, disease resistance and myocardial disease.
It is essential in iodine metabolism and causes iodine deficiency secondarily in the
thyroid gland which is present in all vertebrates, with its hormones playing an
indispensable role in control of the basal metabolic activity. Iodine is essential for
gestational development, particularly of the central nervous system, and deficiency
during pregnancy has negative and irreversible effects on the developing fetus;
postnatal deficiency is associated with cognitive deficits. In ruminants common
problems include abortions, young born dead, weak neonates, increased neonatal
mortality, prolonged gestation and infertility.
Selenium and Iodine Status in Patagonia: concentrations of Se and iodine in most
forages of agricultural systems in southern Chile are deficient, indicating marginal
levels in natural habitat. Levels on the east of the Andes are expected to be even
lower. There are a still focal areas with endemic goiter in Argentinean Patagonia.
Huemul Migrations as a clue to the current predicament?
In autumn, cattle were commonly herded together with huemul to lower elevations
for the winter. Huemul also moved from high Andes down to Pacific coastal areas.
Based on telemetry they were shown to spend the evening and night in valley bottoms
and to move to higher elevations in the morning in valleys not settled or used by
humans. Many huemul lived in bottoms of valleys in Argentina during winters about
60 years ago, though people now residing there have never seen a huemul [2]. In Santa
Cruz, huemul groups of 50 used to migrate annually some 50 kilometers from the high
Andes into the treeless steppe. Others reported huemul at 200 km from the Andes in
treeless grasslands, or wintering groups of 100 huemul 80-100 km from forests,
184 R
resembling migratory behavior of other Odocoilines.
Traditional winter ranges and valley bottoms likely were source areas for huemul
populations and for migratory behavior. Krieg [9] mentioned the existence of all-year
resident huemul in quiet valley bottoms. Migratory huemul in Argentina were likely
eliminated by overhunting, easily being killed as they show no fear. Thousands per
year were killed at rates of 1-2 deer per kmยฒ, coincidently the average density where
they still occur now [2]. Huemul were used to feed people, dogs, chicken and pigs;
their skins were used for shelters for people and domestic animals. With loggers also
killing huemul indiscriminately during the colonization, only the most inaccessible
areas provided refuges for remaining huemul. In 1897, reports based on many Andean
expeditions already mentioned few huemul left due to constant and heavy hunting
pressure. All areas useful as winter ranges for livestock were occupied by settlers early
on or became private, and often were cleared of forests. Migratory behavior is an
acquired trait. Thus, by eliminating the migratory segment of a huemul population, the
remaining animals became tied to a refuge. The few dispersers potentially leaving
such populations would tend to follow prime habitat like valley bottoms and be at high
risk to be eliminated by hunting or dogs.
Discussion
Currently favored reasons for failing huemul recovery in Argentina do not explain
several observations. In healthy populations, losses due to predation by puma, foxes
and feral dogs, and accidents would not present a problem. Domestic and exotic wild
herbivores do not generally affect healthy native cervids elsewhere as most plant-
herbivore systems are multi-species guilds. Domestic and exotic wild herbivores as
disease agents also do not generally affect healthy populations of native cervids, but
undoubtedly increase the risk of introducing epidemiologically important new
diseases. These factors might affect individual huemul, thus current conservation
efforts aim at reducing or eliminating these, but it is not known if such actions have
resulted in improvements. Regardless, populations do not recover even in the absence
of one or several supposedly detrimental factors.
Recruitment in Argentina is too low for recolonizations or numeric increases of
populations. Remaining refuges occur at high elevations near the continental divide,
with high precipitation, far from the Pacific, and in plant communities growing on
soils strongly influenced by igneous rocks, glaciation and volcanism. We posit that
these areas likely provide suboptimal trace mineral levels to huemul. This affects their
reproduction and survival because they lack the opportunity to compensate these
nutritional imbalance by migrating to more favorable sites like valley bottoms and
historical winter ranges.
Deficient dietary iodine and Se levels result in a plethora of problems. Although
ultimate relationships are biochemical, these are expressed in reduced immune
functions, reduced systemic growth and reproductive potential, and behavioral
changes. Thus, the initial diagnosis might be death due to diseases, whereas the
proximate factor might actually be lack of iodine and Se [10]; resistance to diseases
is tied to adequate dietary iodine and Se. Similarly, predation rates might be mistaken
as the problem, whereas the proximate factor might be behavioral changes due to
nutritional deficiency of iodine and/or Se, causing animals to be weak, uncoordinated
and behaviorally affected [10].
R 185
Increasing Se levels in otherwise deficient wild deer increased the recruitment rate
by 260% [11]. Ovis canadensis responded drastically to Se mineral licks [10]. Adults
shed summer coats much earlier, young were weaned much later, and predation losses
diminished. Movement behavior changed once animals had access to Se salt blocks
placed on summer ranges: animals no longer made temporary movements to winter
grounds and valley bottoms to search for natural mineral licks. In contrast, control
sheep without salt blocks continued to make such movements and had lower neonatal
survival. In an additional population, salt blocks on summer ranges were also
provided, but lacking Se; sheep also stopped movements to lower areas, but
recruitment rates were 67% lower than the population receiving Se salt. In
comparison, huemul in Chile also was observed to make summer movements to valley
bottoms (Saucedo, pers. com.), but they would only survive if those areas were free
from people and dog disturbances. The absence of such movement behavior in
Argentine populations likely stems from the continuous elimination of huemul
reaching lower areas populated with people.
Future efforts must be directed beyond the current belief system. The current lack
of science-based monitoring prevents drawing any conclusions about cause and effect.
In the worst case, decisions will be made on unreliable data, thereby compromising
the future of the species. First and foremost is the need to study huemul populations
directly with telemetry as indirect methods will be cost-prohibitive [1]. Specifically,
studies should test our hypothesis.
Reference
[1] Flueck, Smith-Flueck (2006) European Journal of Wildlife Research 52:69-80. [2]
Smith-Flueck 2003. The ecology of huemul in Argentina. Diss., Univ. Nac. Comahue,
Arg.
[3] Flueck (2003) Pp. 30-34 In (Ed.Acosta-Jamett, G.) 4ta reuniรณn Chileno-Argentina
sobre estrategias de conservaciรณn del huemul. CONAF and CODEFF, Las Trancas,
Chile.
[4] Flueck, Jones (2006) Veterinary Parasitology 135:381-383.
[5] Texera 1974. Anales del Instituto de la Patagonia, Punta Arenas (Chile) 5:155-188.
[6] Caughley 1994. Journal of Animal Ecology 63:215-244.
[7] Carter, Robbins, Brown 1970. Journal of Range Management 23:234-238.
[8] Flueck, Smith-Flueck 2006. Wildlife Society Bulletin 34:in press.
[9] Krieg 1940. Als Zoologe in Steppen und Wรคldern Patagoniens. Lehmanns V.,
Germany.
[10] Hnilicka 2001. Biennial Symposium Northern Wild Sheep and Goat Council
13:69-94.
[11] Flueck 1994. Ecology 75:807-812.
(Poster presentation.)
186 R
159
Deer management and private hunting? Turning point for
management system in Japan.
A. Takayanagi
Laboratory of Forest Biology, Graduate School of Agriculture, Kyoto University,
Japan
A new deer management system in Japan will discuss based on historical, cultural and
social background with comparison between other countries.
At the beginning of modern era, similar to European countries, releasing from feudal
system caused exploitation of wildlife by free hunting in Japan. The first hunting law
was established in 1895, 28 years after the restoration. The main purpose of this law
is to regulate hunting to prohibit exploitation or extermination or wildlife. The law did
not function so effective as to prohibit the extinction of wolf. After WWII, the US
occupation administration suggests Japanese government reinforcement of hunting
regulation and the major revision were made in 1963. The number of hunters, which
were increasing rapidly after WWII, have been decreasing after increases of hunting
taxes and fees in the 70's. Deer irruptive increase has come out in several prefectures
in late 80's. Agricultural and forestry damages become huge and decreasing deer
population is urgent issues for many prefectures. Local governments take policy to
increase total hunting bag of deer, but they cannot achieve it because of low hunting
activities. In Japan hunting is not a popular sport. Venison is not popular diet. It is
difficult to pay the management cost by hunting income. Wildlife is not treated as
public goods legislatively as same as Germany. Some people advocate that wildlife
should be stated as public goods in wildlife law but it would be quite difficult because
there is no clear consensus about who pay the managing costs. Though hunters are
decreasing, some farmers get hunting licenses to protect their own cultivated fields.
Local communities have still thought that hunting by the community member may be
more safety than by hunter coming out of the community. Designating deer as local
public goods will provide better management system. In deer management districts,
members can choose public control killing, private hunting by members, and private
hunting by hunter coming from outside of community or mixture of three. Policy
promoting game uses as common property, which has been denied in modern Japan,
will construct next relationships between wildlife and people.
(Poster presentation.)
R 187
160
The biology of antler growth in endangered Bawean deer (Axis
kuhlii).
G. Semiadi
1
, K. Subekti
1
, B. Masyud
2
, I. K. Sutama
3
, and L. Affandy
1
Puslit Biologi LIPI, Gd. Widya Satwaloka Jl. Raya Bogor-Cibinong km.46. Cibinong
16911 Indonesia
2
Fakultas Kehutanan, IPB, Bogor 1600, Indonesia
3
Balitnak Ciawi, Departemen Pertanian, Bogor 1600, Indonesia
Bawean deer (Axis kuhlii) is an endemic and endangered Indonesian deer with their
natural habitat limited to 4500 ha of protected area in Bawean island. Data on their
biology is very limited, similarly to their population status. A study on their biology
of antler growth was conducted for three years in two separate captivities, using
animals from 3-4 months and 22 months old groups. The study showed that pedicle
started to grow at 6 months of age, for 54.7 days, coincide with the testes entering the
scrotum. Spiker velvet antler was noticed to grow 1.5 months later and took 72.3 to
fully-grown. Length of adult stags in hard antler condition was between 88 to 243
days. Hard antlers morphology was relatively small compared to other tropical deer,
where the main beam length was 49% and 25% shorter than sambar and rusa stags,
respectively. The results indicated that the period of antler growth in Bawean deer was
within the normal range of other tropical deer species.
(Poster presentation.)
161
The preservation of rusa stag semen using TRIS egg yolk diluent
with different carbohydrate and storage temperature.
W. M. M. Nalley
1
, R. Handarini
2
, G. Semiadi
3
, M. R. Toelihere
4
, T. L. Yusuf
4
, and
B. Purwantara
4
1
Fakultas Peternakan, Universitas Nusa Cendana, Kupang, Nusa Tenggara Timur,
Indonesia
2
Fakultas Pertanian, Universitas Sumatra Utara, Medan, Indonesia
3
Pusat Penelitian Biologi LIPI, JL. Ir. H.Djuanda 18, Bogor 16002, Indonesia
4
Departemen Reproduksi dan Kebidanan, Fakultas Kedoteran Hewan, Institut
Pertanian Bobog, Bogor, Indonesia
The purpose of this research was to study the viability of the rusa stags (Cervus
timorensis) semen preserved at 27-28 C and 35 C in tris extender and supplemented
with various sources of carbohydrate, i.e. glucose (TG), fructose (TF), and sucrose
(TS). Collection of semen was conducted from five adult stags at the hard antler
188 R
condition. Semen was collected with electro ejaculator after sedation of the stags using
a combination of 1 mg xylazine and 2 mg ketamin i.m. kg-1 body weight. Semen was
evaluated by macroscopic and microscopic methods, with the concentration of 100
million motile sperm ml-1 were stored and observed every 3 hours (27-28 C) and 24
hours (35 C). Data were analysed with completely random design, and means were
compared by the least significant difference test. The collected fresh semen had a
volume of 2.06 ml, pH 7.03, yellow white until cream in color and its consistency
ranged from normal to thick. Mass movement was ++ to +++ and percentage of motile
spermatozoa (MS) 75.83%, percentages of live spermatozoa (LS) 87.67%, abnormal
spermatozoa (NS) 7.31%, intact plasma membrane (IPM) 76.83%, and intact
acrosome cap (IAC) 80.17%, with concentration of spermatozoa was 842.35 x 106 ml-
1.
At 27-28 C preservation temperature and up to 15 hours of the observations, the
glucose treatment (TG) showed a significantly higher (p<0.05) in any parameters
being evaluated (MS 43.50%, LS 52.50%, IPM 47.67%, IAC 51.50%) than that the
fructose (TF) or sucrose (TS) treatments (42.50%; 45.17%; 41.00%; 41.50% vs.
42.50%; 46;00%; 41.33%; 43.33%, respectively). The motility of sperm which was
down to 42.50% motile was reached at 12 hours in both TF and TS treatments. At 35
C preservation temperature, the TG and TS showed a significantly different (p<0.05)
in all parameters being evaluated (MS 42.22%, LS 53.89%, IPM 45.56%, IAC 47.67%
and MS 42.67%, LS 48.89%, IPM 45.67%, IAC 46.22%, respectively) than in the TF
treatment (MS 33.33%, LS 40.89%, IPM 36.44%, IAC 38.78%). From this study, it
was concluded that the preserved rusa stag semen can be used for artificial
insemination purposes up to 5 hours post preservation at 27-28 C using TG as
carbohydrate sources, while in TS and TF carbohydrate sources were 12 hours. Semen
preserved at 3-5 C, in TG and TS can be stored effectively up to nine days, whereas
in TF was only for six days.
(Poster presentation.)
R 189
162
Semen quality of rusa stags (Cervus timorensis) during one antler
cycle.
R. Handarini
1
, W. M. M. Nalley
2
, G. Semiadi
3
, M. R. Toelihere
4
, S. Agungpriyono
4
,
B. Purwantara
4
, and Subandriyo
5
1
Fakultas Pertanian, Universitas Sumatra Utara, Medan, Indonesia
2
Fakultas Peternakan, Universitas Nusa cendana, Kupang, Nusa Tenggara Timur,
Indonesia
3
Pusat Penelitian Biologi LIPI, JL. Ir. H.Djuanda 18, Bogor 16002, Indonesia
4
Departemen Reproduksi dan kebidanan, Fakultas Kedoteran Hewan, IPB, Bogor,
Indonesia
5
Balitnak, Pusat Penelitian Peternakan, Departemen Pertanian, Ciawi Bogor,
Indonesia
Circanual pattern of the antler cycle correlated with the circanual pattern of testicular
function and reflecting changes in semen production. The objective of the research on
the reproductive biology of rusa stags (Cervus timorensis) was to quantify the semen
quality related to the their natural antler cycle, i.e. casting, velvet and hard antler
stages. The research was conducted for 18 months in four adult rusa stags (4 to 6 years
old). Data were collected at three weeks intervals during the annual antler cycle and
semen characteristics were evaluated macroscopically and microscopically. The
results indicated that semen quality was significantly lower (P<0.05) at velvet antler
stage compared to that at hard antler stage (mass movement 0.63 vs. 2.38, motility
32.5% vs. 72.76%, concentrations 199.56 vs. 1055.95 million/ml, live spermatozoa
50.87 vs. 79.40%, abnormality 35.59 vs. 9.80%, respectively). The nutritional quality
of seminal plasma in velvet vs. hard antler conditions showed a significant difference
(p<0.05) in all parameters being observed (3.13 vs. 25.93 calorie, ash content 130 vs.
15140 mg/100 ml, protein 134 vs. 4560 mg/100 ml, fat 204 vs. 212 mg/100 ml, Ca 3.6
vs. 6.5 mg/100 ml, respectively). It was concluded that the best semen quality was
produced and obtained at hard antler stage, which was also as the longest stage of the
antler cycle, starting from June to February.
(Poster presentation.)
190 R
Problems of deer
overabundance
192 R
163
A test of localized management in a white-tailed deer herd.
B. F. Miller
1
, R. W. DeYoung
2
, T. A. Campbell
3
, B. R. Laseter
1
, W. M. Ford
4
, and K.
V. Miller
1
1
Warnell School of Forest and Natural Resources, University of Georgia, Athens, GA;
USA
2
Caesar Kleberg Wildlife Research Institute, Texas A&M University-Kingsville,
Kingsville, TX; USA
3
USDA/APHIS/WS/NWRC-Texas Field Station Texas A&M University-Kingsville,
Kingsville, TX, USA
4
USDA Forest Service, Northeastern Research Station, Parsons, WV, USA
Herbivory by white-tailed deer within forest regeneration areas can have profound
impacts on stand structure, composition, and biodiversity. Because traditional
management strategies (i.e. sport hunting) are not solving the damage problem in
many areas, localized management has been proposed as a possible solution.
Localized management involves the โsurgicalโ removal of one or more matriarchal
social groups in an area deemed sensitive to browsing pressure. Because white-tailed
deer females are generally philopatric to their natal area, removal of entire social
groups may create low-density zones for >5 years. However, this technique has only
been tested in a migratory, low-density, and un-hunted deer herd in New York. In that
study, deer did not recolonize areas previously occupied for > 2 years after the
removal of an entire social unit. Herein, we describe our experimental investigation
of localized management in a high-density and non-migratory white-tailed deer herd
in the Appalachian Mountains of West Virginia, USA. We used a combination of
radiotelemetry, sightings of tagged animals, and genetic data to: 1) investigate the
movement ecology and social structures of white-tailed deer, 2) design and conduct
an experimental removal of a social group(s) in the vicinity of a selected forest
regeneration area(s), 3) monitor movements of adjacent social groups to detect
encroachment or dispersal/colonization into the removal area, and 4) determine the
relationship of colonizers (both marked and unmarked) to social groups and marked
individuals. Between 24 February 1999 and 19 March 2005 we captured 358
individual deer. We collected DNA samples, ear-tagged each animal, and radio-
outfitted 275 of the animals. Animals were located 3 times weekly from permanent
geo-referenced triangulation stations. A total of 47,282 radio-telemetry locations were
obtained between 7 April 1999 and 27 April 2005. We recorded 17,731 visual
observations of tagged and non-tagged deer from April 1999 to April 2005. A 1.2 km
2
area was established as an experimental removal area based on social groups
delineated via radio-telemetry. The experimental removal was conducted from 7
January to 27 February 2002 and accomplished by trapping and sharpshooting. A total
of 51 deer were removed during the seven week removal period. Track counts in the
snow conducted before, during, and after the removal indicated that the removal was
largely successful. A final removal of deer was conducted from 7 January to 21
February 2005 to determine if animals subsequently occupying the removal area were
R 193
survivors of the initial removal, their descendants, or immigrants. A total of 31 deer
were removed during the second removal. DNA was extracted from genetic samples
using commercially available kits, and amplified through polymerase chain reaction.
An ABI Genescan 3130 sequencer was used to examine allele sizes for the panel of
14 polymorphic microsatellite markers. A spatial autocorrelation analyses based on
pairwise Moranโs I among 229 individual adult (>1.5 yrs.) females revealed that
genetic relatedness was inversely related to the distances between core areas
determined by telemetry data or trapping location. Females with core areas separated
by less than 1km were more related than expected by random chance. However,
autocorrelation became non-significant at distances >1km and declined to 0.0 at
2.1km. The correlogram displayed a stabilizing pattern, suggesting spatial structure
at the group level. Using visual observations we were able to delineate 28 social
groups that contained at least two members with genetic information. Mean
relatedness within groups was 0.1, a value similar to that of first cousins. This
evidence of fine-scale social group structuring indicates that the theoretical basis of
localized management applies on the study site; females were indeed philopatric to
their natal area and were structured in social groups containing related individuals.
However, despite evidence for social group structuring of the deer herd, localized
management was ineffective in establishing an area of reduced deer density on this
study site. When we compared the genetic data collected from the first and second
removals, we found evidence of genetic differentiation (significant Fst, and exact tests
of differentiation, frequency of private alleles, and Hardy-Weinberg disequilibrium).
The large number of animals removed three years after the initial experiment and the
evidence of genetic differentiation between the first and second removal groups
indicates recolonization by immigrant deer. We therefore conclude that localized
management may not be applicable in areas of high deer density.
(Oral presentation.)
164
Do wildlife warning reflectors alter white-tailed deer behavior along
roadways?
G. J. D'Angelo
1
, J. G. D'Angelo
1
, G. R. Gallagher
2
, D. A. Osborn
1
, K. V. Miller
1
, and
R. J. Warren
1
1
University of Georgia, Athens, GA; USA
2
Berry College, GA; USA
Deer-vehicle collisions are increasing in frequency throughout much of the United
States. Government transportation agencies seek to minimize deer-vehicle collisions
through a variety of techniques. However, few mitigation strategies have undergone
extensive independent testing of their effectiveness prior to deployment in the field.
Wildlife warning reflectors are marketed as an effective and humane technique for
reducing wildlife-vehicle collisions. Yet, previous studies have provided a limited
194 R
understanding of reflector efficacy. We evaluated the effectiveness of 4 colors of
wildlife warning reflectors (red, white, blue-green, and amber) for altering deer
behavior that might help prevent deer-vehicle collisions. We observed white-tailed
deer (Odocoileus virginianus) behaviors relative to roads before and after installation
of wildlife warning reflectors using a forward looking infrared camera. From 18
November 2004-1 May 2005, we recorded 1,370 deer responses to vehicles during 90
nights of observations (4-hr each). Our study contradicted claims by the manufacturer
that wildlife warning reflectors deter deer from crossing the highway when reflecting
vehicle headlights. Our results demonstrated that deer exposed to each of the 4 colors
of reflectors we tested were more likely to be involved in negative deer-vehicle
interactions than without the devices present. We concluded that wildlife warning
reflectors were ineffective in changing deer behavior such that deer-vehicle collisions
might be prevented.
(Oral presentation.)
165
Cascading effects of long term chronic browsing on lifehistory traits
in white-tailed deer.
S. D. Cรดtรฉ, A. Simard, R. B. Weladji, and J. Huot
Department of Biology and Centre d'etudes Nordiques Laval University, Quรฉbec,
Canada
The recent increase in deer populations worldwide is negatively affecting vegetation
in many areas, but few studies have assessed the long-term cascading effects on the
life histories of Cervids. White-tailed deer were introduced on Anticosti Island
(Quรฉbec, Canada) in 1896 and expanded rapidly so that evidence on vegetation of
severe deer browsing was noticed by the early 1930s. Detailed monitoring of
vegetation and deer life histories, however, only started in the mid 1970s. Here, we
assessed whether chronic browsing contributed to a decline in the quality of deer
summer/autumn diet during the last 25 years, by comparing percentage of nitrogen of
rumen contents between 1977-79 and 2002-04. We further assessed the cascading
impacts on deer body condition and reproduction. Rumen nitrogen content declined
22% between 1977-79 and 2002-04, suggesting a reduction in forage quality in late
summere-arly autumn. We also found a decline in asymptotic autumn body mass for
both males and females, and a light decline in foot length for males. When controlling
for annual stochasticity, age and date, males were about 9% smaller and also tended
to accumulate mass more slowly in 2002-04 than in 1977-79. Similarly, at the end of
November, adult does were 6% smaller in recent years than in 1977-79. The
probability of ovulation increased 12% between the two periods, but twinning rate
declined 7% resulting in a higher ovulation rate (9%) in 2002-04, although not quite
significant. Our results suggest that following a persistent decline in forage quality
over a period of 25 years white-tailed deer have adjusted their life history traits to
R 195
maintain reproduction but at the expenses of growth.
(Oral presentation.)
166
Regeneration dynamics of boreal forests along an experimental
gradient of deer densities.
J.-P. Tremblay
1,2
,J. Huot
1,2
and F. Potvin
1,3
1
Chaire de recherche industrielle CRSNG-Produits forestiers Anticosti, Dรฉpartement
de biologie, Universitรฉ Laval, Quรฉbec, Canada
2
Centre dรฉtudes nordiques, Universitรฉ Laval, Quรฉbec, Canada
3
Ministรจre des Ressources naturelles et de la faune, direction de la recherche sur la
faune, Quรฉbec, Canada
Deer are key components of forest ecosystems where they act as a chronic disturbance
modulated by their density. Loosely regulated deer population can alter plant
community structure and composition. This is especially critical for late successional
forests that are vulnerable to compositional shifts following events that suppress
advance regeneration. Although impacts are generally assumed to be proportional to
deer density, nonlinearities may emerge e.g. from deer induced modifications to
competitive interactions between plants. As a case study, we investigated the
relationships between the regeneration dynamics of eastern balsam fir Abies balsamea
forests and white-tailed deer Odocoileus virginianus density on Anticosti Island, Ca.
We hypothesized that the responses of the field layer plants and advance tree
regeneration are either: (1) directly proportional to deer density; (2) a smooth
nonlinear function of deer density; or (3) a nonlinear function with thresholds. We
tested predictions from these hypotheses by experimentally manipulating both deer
densities (0, 7.5, 15, 27 and 56 deer/km
-2
) and forest cover. In clear-cuts, the dominant
responses of the field layer plants were growth and reproduction suppression at high
deer densities and exponential recovery at densities ~<15 deer/km
-2
. The mortality rate
of balsam fir seedlings increased exponentially with deer densities, with the
cumulative mortality reaching 74 8% at 56 deer/km
-2
. Browse tolerant species such as
grasses were positively and nonlinearly related to deer density suggesting an apparent
competitive gain. Similarly, the abundance of spruce Picea spp. saplings was unrelated
to deer density and increased with time as seedlings were recruited into larger height
classes. Such rapid changes in the early successional stages have long term indirect
consequences on successional patterns through processes such as seed availability,
germination and early establishment of seedlings. Selective browsing at high deer
densities over an extended period of time thus sets the conditions for recruitment
failure following a stand-replacing disturbance. We propose a conceptual model based
on the catastrophe theory to predict the evolution of deer- forest systems. On
Anticosti, reduction of local deer densities to levels <15-7.5 deer/km
-2
in the first three
years following a clear-cut appears compatible with the regeneration of native forests.
196 R
(Oral presentation.)
167
Impacts of cervids on invertebrate communities on forest floor in
relation to deer species, density and site productivity.
O. Suominen
1
, I. L. Persson
2
, and T. Saikkonen
1
1
Section of Ecology, Dept. of Biology, Univ. of Turku, Finland
2
Dept. of Animal Ecology, Swedish Univ. of Agricultural Sciences, Umeรฅ, Sweden
The impacts of high ungulate densities on vegetation have been amply studied and the
impacts on some popular animal groups, such as birds, have even awoken public
concern. The potential impacts of cervids on invertebrates have received much less
attention. We studied how moose and reindeer at different densities and seasons affect
vegetation and invertebrate fauna on the floor of boreal forest. We simulated
browsing, urinating, and faecal deposition corresponding to 4 different moose
densities in 8 exclosures along a forest productivity gradient in Sweden. Our results
show that moose at higher population densities can have a substantial impact on the
ecosystem, and that the effect can be modified by habitat productivity: As a result of
these changes both vegetation and vegetation living invertebrate assemblages changed.
Richness responses of spiders to browsing were negative at unproductive habitats
whereas in the most productive sites moose had a positive impact. The impacts of
reindeer grazing were studied comparing heavily grazed Finnish side and ungrazed
Russian side of the reindeer fence on the Finnish-Russian border in Lapland.
Vegetation was lower, temperature was higher and humidity lower in grazed plots.
Soil moisture was higher in ungrazed plots under thick Cladina-lichen cover in winter
ranges. In these pine forests we found spider species characterizing boggy habitats
from the ungrazed plots and species favouring dry open habitats from the grazed plots.
The relationship between site productivity and reindeer impact on invertebrate
richness was opposite to that observed for moose: The negative impact of reindeer on
spider and beetle richness increased with increasing productivity. The impact on
spider abundance and richness differed between moose and reindeer, but in both the
impact of browsing/grazing on spiders correlated with the impact of cervids on the
abundance of the prey of the spiders.
(Oral presentation.)
R 197
168
Sustainable population density of red deer in Mediterranean
ecosystems.
J. Carranza, J. Torres, S. Alarcos, J. Pรฉrez-Gonzรกlez, C. B. Sรกnchez-Prieto, C. Mateos,
L. Castillo, and J. Valencia
Biology & Ethology, University of Extremadura, Cรกceres. Spain.
How population density translates into overabundance is a complex issue that depends
on ecological conditions. Previous information suggests that red deer (Cervus elaphus)
populations in Mediterranean habitats of Southwestern Spain may reach higher
densities than in any other parts of Europe. Our goal in this study was (1) to describe
the densities occuring in main areas of red deer distribution in Southwestern Spain,
and (2) to explore the relationships between these densities and some indicators of
performance and sustainability. We found that population density in hunting estates
ranged from 0.1 to 1.0 individuals per hectare, averaging between 0.29 and 0.39
individuals per hectare in three main large regions included in the distribution area.
We explored body size of males and females, antler size and fertility for individuals
of different ages, as well as impacts on arbustive vegetation, as biological indicators
of overabundance. All these parameters related to density, and younger individuals
appeared more sensitive to population density. However, in our populations, these
biological indicators were only weakly affected by increases in density compared with
the case for other European populations. Two main reasons may contribute to explain
this difference. First, productivity of Mediterranean ecosystems for wild herbivores
is very high, which also corresponds with high levels of biodiversity, and may explain
why red deer use so small home ranges in these types of habitats compared to other
areas of Europe. Second, natural vegetation can buffer the limiting season in
Mediterranean habitats (schrub plants during the summer) better than in northern
latitudes, where cold winters may set a very low level for carrying capacity. Moderate
densities constitute an appropriate reccomendation for management that can put
together objectives of trophy quality and habitat conservation, but the posibility that
sustainable densities in Mediterranean areas may be higher than in other parts of
Europe, opens up an interesting oportunity to promote the exploitation of wild
ungulates in these areas.
(Poster presentation.)
198 R
169
Influence of population density on white-tailed deer foraging
behavior and activity budget.
M.-L. Coulombe, S. D. Cรดtรฉ, and J. Huot
Dรฉpartement de biologie and Centre dโรฉtudes nordiques, Universitรฉ Laval, Quรฉbec
G1K 7P4, Canada
At the end of the 19
th
century, 220 white-tailed deer (Odocoileus virginianus) were
introduced on Anticosti Island (Quรฉbec, Canada), and in the absence of predators the
population irrupted. Vegetation communities considerably changed because of the
impacts of severe browsing. Generally, available resources per individual decrease
when population density increases. To assess the effects of high population density on
the foraging behavior of deer in low plant biomass communities, we experimentally
manipulated deer density. We used three sites where forest was partially harvested.
Each site was composed of 2 experimentally-controlled densities (7.5 deer/km
2
and
15 deer/km
2
) in large enclosures, and a natural site where density was estimated at 25
deer/km
2
. At each site-density, three deer were equipped with conventional
radiotelemetry collars and were located a total of 2936 times by radio-tracking.
Vegetation sampling allowed us to compare how deer use available space in relation
to deer density and plant biomass. Individuals used clear-cuts and forested areas in a
similar proportion. Activity budgets were also continuously obtained using motion
sensors and an automatic receiver-datalogger. We compared hourly movements and
activity of deer between density treatments. Adults were less active than yearlings at
7.5 but not at 15 deer/km
2
. Otherwise, movements, activity budgets and available
biomass were similar between 7.5 deer/km
2
and 15 deer/km
2
. However, the available
biomass increased through years and activity budgets varied accordingly. As biomass
increased, deer increased the number of daily activity bouts, which also became
shorter. Deer moved more at dawn and dusk than at other periods of the day. Females
moved less at high densities than at low densities, but not males. This study
demonstrates that relationships between density and foraging behavior are complex
and that controlled-density experiments may help to understand the behavior of
herbivores in relation to available resources.
(Poster presentation.)
R 199
170
Trade-off between food and cover: summer movements and activity
budget in white-tailed deer.
A. Massรฉ, S. D. Cรดtรฉ, and J. Huot
Dรฉpartement de biologie and Centre dโรฉtudes nordiques, Universitรฉ Laval, Quรฉbec
G1K 7P4, Canada
Habitat selection in ungulates is frequently viewed as the result of a trade-off between
food and cover. During summer, individuals are expected to trade-off forage
abundance and escape or thermal cover. White-tailed deer-habitat relationships on
Anticosti Island (Quรฉbec, Canada) are unique because of the absence of predators and
the high deer density (> 20 deer/km
2
locally) in a boreal environment submitted to
severe climatic conditions in winter. Our objectives were to determine the relationship
between habitat use and deer behavior (foraging or resting) and to assess the influence
of forage abundance on movements and activity budget of deer under natural
conditions. We evaluated habitat selection and deer behavior using GPS collars
equipped with activity sensors from July to November (n=19). We sampled vegetation
in clear-cuts, peatlands and forests in deer home ranges to assess forage abundance
within each habitat. Our results showed that habitat selection was influenced by type
of activity: deer foraging preferred open habitats such as clear-cuts and peatlands,
while deer resting increased their use of forested habitats, except at dusk. Spatial
variation of forage abundance influenced deer movement rates. When forage
abundance was high, i.e. in clear-cuts and peatlands, deer moved less than in forested
habitats, suggesting that searching time for forage was reduced. Interestingly, deer
moved rapidly when crossing edges between open and forested habitats, maybe
because they were relocating. Temporal variation of forage abundance and quality
throughout summer did not, however, affect movement rates and activity budget.
Forage abundance was the main factor influencing habitat selection when deer were
foraging, but while resting deer selected more forested habitats.
(Poster presentation.)
200 R
171
Relationships between moose (Alces alces) pellet groups and
characteristics of forests.
R. Heikkilรค
Finnish Forest Research Institute, Vantaa Research Centre, Box 18, FI-01301 Finland
Introduction
Contradictory opinions of moose population size and of methods to estimate
population density as well as browsing pressure indicate needs of more accurate
information for management purposes. Counting fecal pellet group has for long been
analyzed in order to determine habitat use and population density of moose and deer
(Neff 1968, Franzmann et. al. 1976). The pellet group method is considered important
for smaller deer species which are difficult to observe (Cederlund & Liberg 1995).
Winter conditions often restrict seriously aerial census of moose populations and
counting of pellet groups is easier for moose owing to better visibility of pellets in
varying field conditions. Comparison with aerial counting has shown that reliable
information of moose density is possible using pellet group counting (Jordan et al.
1993). However, the important basic factors like behavioral aspects and diurnal
defecation rate (Oldemayer & Franzmann 1981) may vary considerably depending on
region and habitat quality (Andersen et. al. 1992). Besides indicating behavior,
distribution and density of animals, the pellet groups can give related information of
the occurrence and influence of browsing on forests (Hรคrkรถnen & Heikkilรค 1999).
The aim of the study is to determine the characteristics of forest stands and moose
browsing effects correlating with the occurrence of pellet groups in managed forest
area. Special attention is given to possibilities to obtain more information of the
relationships between browsing pressure and moose density.
Material and methods
Field inventory was done in 2004 in four different forest areas in Central Finland
in early summer, when the disappearance of pellets was not supposed to be problem
(Persson 2003). In total 22 lines (92,75 km) were inspected. In Kiltua and Liperi the
lines were situated at 0.5 km intervals, and in Lakomรคki and Jokimaa the study was
done on game triangles (3 x 4 km each), that are yearly used for monitoring game
species in Finland (cf. Linden et.al. 1996),. Measurements were done on 50 m
2
plots
at 50 m intervals. Following information was collected: number of new (previous
winter) and old pellet groups, forest site type, forest stand characteristics (regeneration
area, small seedling stand, sapling stand < 4 m, advanced sapling stand, young timber
stand, advanced timber stand, old timber stand, open or seed tree stand), main tree
species, effect of browsing on different tree species (% removed of foliage). In
addition, the height of young trees up to 6 m was measured in height classes of 1 m.
Counting the pellet groups was done in early summer, as they could be easily
determined before vegetation cover. The new groups dropped in the previous winter
were separated from the old ones. Determining was done according to the differences
in the position of pellets in relation to the ground vegetation and/or their deformations.
R 201
Those pellet groups of which the main part was inside the plot were included.
Calculation to estimate moose density was done using the formula (D/A)(TxF), where
D is the number of pellet groups, A the total area sampled, T the days of pellet groups
accumulation, and F the defecation rate per day and individual (Olsson 1997). As the
accumulation period was used 250 days, and as defecation rates 21 and 24 pellet
groups per day were compared (cf. Jordan et.al. 1993, Andersen et. al. 1992).
Mortality during accumulation period was not included in calculation.
Statistical comparisons were done using the Pearson correlation and one way
ANOVA programs of the SPSS-procedure.
Results
Moose pellet group density varied significantly between the stand age classes in
three study areas (p <0.001, one way ANOVA). 60-75 % of pellet groups were found
in the seedling stands and advanced young forest stands. The Liperi study area
differed somewhat with no respective correlation between stand classes. In that more
spruce dominated area the middle-aged, young timber stands also had relatively high
numbers of pellet groups. In Liperi the forest peatland areas had on average more
pellet groups than the mineral soil forests (p < 0.05), contrarily to the other areas with
more groups in the latter kind of forests (p < 0.001 for each area).
The fecal output of moose in winter had most consistently occurred close to young
Scots pines (Pinus sylvestris). Both the number of pines and the effect of moose
browsing in terms of damage caused to the tops of pines correlated well with pellet
groups (Table 1). Both silver birch (Betula pendula) and downy birch (Betula
pubescens) appeared also to be often used by moose in the close vicinity of pellet
groups. In Lakomรคki and Liperi areas even overbrowsing of pines by moose correlated
significantly with pellet groups. Overbrowsing of both birch species was consistently
correlated with pellet groups in all study areas excluding Lakomรคki, where only
downy birch correlated. These results obviously reflect also browsing pressure
depending on food availability in tree species.
Table 1. Characteristic of forests correlating with the number of moose pellet groups
in four study areas in Central Finland. Statistical significance * = 0.05, ** = 0.01.
x = browsing data is included in "top damage".
Characteristics
measured on
28 study lines
Kiltua Liperi Jokimaa Lakomรคki
% of lines with
correlation to
pellet groups
No. of pine 0.142** 0.327** 0.375*
*
0.203** 50
No. of silver
birch
0.093* 0.220*
*
11
No. of downy
birch
0.120** 0.323** 29
202 R
Characteristics
measured on
28 study lines
Kiltua Liperi Jokimaa Lakomรคki
% of lines with
correlation to
pellet groups
No. of aspen 0.167*
*
7
No. of juniper 0.073* 4
Height of
young pines
0.190** 7
Height of
young spruces
-0.134* 4
Height of
young D.
birches
-0.145** -0.229** 7
Browsing on
pine
0.272** 0.287** x x 39
Browsing
on S. birch
0.230** 0.116** x x 21
Browsing
on D. birch
0.338** 0.123** x x 25
Browsing
on willows
0.078* 7
Browsing
on juniper
-0.105** 7
Top damage
of pine
0.245** 0.351** 0.345*
*
0.277** 61
Top damage
of S. birch
0.239*
*
7
Overbrowsing
of pine
0.188** 0.282** 14
Overbrowsing
of S. birch
0.240** 0.154** 0.183*
*
21
Overbrowsing
of D. birch
0.256** 0.146** 0.142* 0.356** 25
R 203
Characteristics
measured on
28 study lines
Kiltua Liperi Jokimaa Lakomรคki
% of lines with
correlation to
pellet groups
Overbrowsing
of aspen
0.073* 0.156*
*
11
Condition of
pine
-0.191** -0.266** 7
Condition
of D. birch
-0.273*
*
4
The deciduous tree species known to be selected by moose were not commonly
correlating with pellet groups. Aspen (Populus tremula) appears to be to some extent
associated with pellet groups, but willows (Salix spp) only rarely and for instance
rowan (Sorbus aucuparia) is lacking. It can be noticed that aspen is included as
overbrowsed by moose.
According to the numbers of pellet groups moose density estimations in the study
areas were between 1.7 and 3.2 moose/km
2
with daily output of 21 groups, and
between 1.5 and 2.8 moose/km
2
with the output of 24 groups per day. The highest
moose density was calculated in Lakomรคki and the lowest in Kiltua and Liperi.
Discussion
Pellet group counts have mostly been used in habitat selection analyses (Cairns &
Telfer 1980) as well as for comparison between aerial counting in order to find
alternatives to accurate moose density estimation (Jordan et. al. 1993). In the present
study pellet groups were used to obtain related information of the forest stands under
the effect of browsing. Analyzing pellet groups and habitat preferences include
uncertainties i.e. in time spent by animals (Guillet et. al.1995). On the other hand,
correlation has been found between the occurrence of pellet groups and deer bedding
sites (Collins & Urness 1981). Studies of moose browsing show that pellet group
density correlates significantly with the amount of browsing in young pine stands
(Heikkilรค & Hรคrkรถnen 1993).
The lack of correlation between pellet groups and certain tree species can be
explain simply by their reduced number in the studied forest areas. In the moose
winter ranges population density and consequently browsing pressure on selected tree
species can be relatively high. The effect of browsing depends also on the available
food resources. Considerable difference was found for instance between the Liperi and
Lakomรคki study areas. The former one is more spruce dominated than the latter one,
which as relatively dry forest is typically dominated by pine forests. Estimating moose
population density with pellet group counts includes uncertainty factors such as
defecation rate in different habitats (Andersen et. al. 1992) and accurate timing of
defecation (Persson 2003). However, in the present study the comparison between
areas suggests that at similar moose density the effects of browsing can differ
indicating difference also in the sustainability of forest to browsing.
Determining more accurately relationships between moose population size and
204 R
browsing damage is a relevant management question. In Finland suggestions have
been made of using pellet group counts for population density estimates in the wildlife
triangle network (Tiainen 1998). Planning an efficient sampling method for pellet
group counts also other possibilities than triangle sampling could be taken into
account, including monitoring the effects of moose browsing pressure.
References
Andersen, R., Hjeljord, O. & Saether, B-E. (1992) Moose defecatio rate in relation to
habitat quality. Alces 28:95-100.
Cairns, A.L. & Telfer, E.S. (1980) Habitat use by 4 sympatric ungulates in boreal
mixedwood forest. J. Wildl. Manage 44:849-857.
Collins, W.B. & Urness, P.J. (1981) Habitat preferences of mule deer as rated by
pellet-group distributions. J. Wildl. Manage. 45(4):969-972.
Franzmann, A.W., Arneson, P.D. & Oldemayer, J.L. (1976) Daily winter pellet groups
and beds of Alaskan moose. J. Wildl. Manage. 40:374-375.
Guillet, C., Bergstrรถm, R., Cederlund, G., Bergstrรถm, J. & Ballon, P. (1995)
Comparison of telemetry and pelet-group counts for determining habitat selectivity by
roe deer (Capreolus capreolus) in winter.
Heikkilรค, R. & Hรคrkรถnen, S. (1993) Moose (Alces alces L.) browsing in young Scots
pine stands in relation to the characteristics of their winter habitats. Silva Fennica
27:127-143.
Hรคrkรถnen, S. & Heikkilรค, R. (1999) Use of pellet group counts in determining density
and habitat use of moose Alces alces in Finland. Wildl. Biol. 5:233-239.
Linden, H., Helle, E., Helle, P. & Wikman, P. (1996) Wildlife triangel scheme in
Finland: methods and aims for monitoring wildlife populations. Finnish Game
Research 49:4-11.
Neff, D.J. (1968) The pellet group count technique for big game trend, census, and
distribution: a review. J. Wildl. Manage. 32:597-614.
Oldemeyer, J.L. & Franzmann, A.W. (1981) Estimating winter defecation rates for
moose, Alces alces. Canadian Field-Naturalist 95:208-209.
Olsson, O., Wirtberg, J., Andersson, M. & Wirtberg, I. (1997) Wolf (Canis lupus)
predation on moose Alces alces and roe deer Capreolus capreolus in south-central
Scandinavia. Wildlife Biology 3:13-25.
Persson, I-L. (2003) Moose population density and habitat productivity as drivers of
ecosystem processes in northern boreal forests. Acta Universitatis Agriculturae
Suecicae Silvestria 272. 30 p.
R 205
Tiainen, J. (1998) Miten valkohรคntรคpeuran ja metsรคkauriin runsauden seuranta tulisi
jรคrjestรครค? (In Finnish with English summary: Organization of small cervid monitoring
in Finland). Suomen Riista 44:37-42.
(Poster presentation.)
206 R
Conservation of free
ranging populations:
conflicts of interest
208 R
172
Status, distribution and conservation of musk deer (Moschus
chrysogator) in Kedarnath Wildlife Sanctuary, Uttranchal
Himalayas, India.
O. Ilyas
Department of Wildlife Sciences, Aligarh Muslim University, Aligarh-202002, India
Out of the four species of musk deer i.e. Himalayan musk deer (Moschus chrysogaster
Hodgson), Siberian musk deer Moschus moschiferus Linnaeus), Dwarf Musk deer
(Moschus berezovskii Flerov) and the Black musk deer (Moschus fuscus Linnaeus),
only one species Moschus moschiferous is present in Himalayan range of India. The
musk deer has been exploited for the musk a secretion of the preputial gland of the
male, which is used in perfumery and medicine for centuries. The musk been very
expensive animal product fetching up to US$ 65000/kg in the international market is
clear inducement to the human greed to put enormous pressure on the animal pursuing
them towards the extinction. The species is listed as โVulnerableโ by IUCN (1974) in
Red data book. Similarly the Wildlife Protection Act (1972) of India put them under
Schedule I species. The first ever-ecological study was done by Green (1985) and later
by Satyakumar (1994) in Kedarnath Wildlife Sanctuary and this study (Ilyas, 2004).
However it is highly desirable that status, distribution, and threats to conservation of
musk deer should be given due importance. To understand the status distribution and
conservation status an intensive survey was conducted at three sites of Kedarnath
Wildlife Sanctuary during premonsoon 2004, between the altitudinal range from
2500m at sea level to 4000 m a.s.l. The direct as well indirect methods were used and
found that the pellet group density of musk deer was found to be maximum in
Saukhark (53.14ยฑ9.6) followed by Tungnath (14.86ยฑ8.71) and Madhmaheshwer
(10.19ยฑ3.2). Principal Component Analysis (PCA) was performed to understand the
habitat use of musk deer. Due to illegal poaching and hunting the population of musk
deer has declined and now confined small fragmented patches and needs urgent
attention for the conservation.
(Oral presentation.)
R 209
173
Seed dispersal by the reintroduced Persian fallow deer in the
Judean Mountains, Israel.
R. Zidon
1
, D. Saltz
2
, and U. Motro
3
1
Environmental Science Program, The Hebrew University of Jerusalem, Israel
2
Mitrani Department of Desert Ecology, Blaustein Institute for Desert Research, Ben
Gurion University, Israel
3
Department of Evolution, Systematics and Ecology, The Hebrew University of
Jerusalem, Israel
The introduction of an animal to a new environment can have significant effects on
the plants population density and distribution. The Persian fallow deer (Dama dama
mesopotamica) has been gradually reintroduced in Israel since 1996. Here we report
a recent study on the potential of the Persian fallow deer as a seed disperser via
endozoochory, and the influence of the deer ingestion on tree seeds. 160 samples of
faeces were collected from the Judean Mountains, near Jerusalem, over a one year
period. Each sample of feces was divided into three parts: the contents of one third
were analyzed in the lab, and the seeds were identified and counted. Another third of
each sample was sown in a greenhouse, and the viability of the seeds was determined
by the proportion which germinated. The last third of the sample was sown in the
field, to test if the process occurs in nature as well. More than 30 species of herbs
germinated from the feces, and germination was stronger (both in the number of
species and in the proportion of germinating seeds) in the greenhouse than in the field.
The species that germinated were from all range of habitat types used by the deer. Of
the trees, carob (Ceratonia siliqua) seeds were the only intact seeds found in our feces
samples. The ingestion had a significant positive effect on the carob seeds compared
to seeds either from intact carob pods, exposed carob seeds, or carob seeds that were
collected beneath trees, and even from carob seeds that were collected from jackal
feces. Our results provide evidence to the great influence that the reintroduced Persian
fallow deer may have on the local Mediterranean flora by contributing to the long-
distance seed dispersal mechanism and specifically to the distribution of carob trees.
(Oral presentation.)
210 R
174
Deer management and monitoring of browsing impacts in Austrian
national parks.
R. Zink and F. Reimoser
Research Institute of Wildlife Ecology, Vienna Veterinary University, Vienna, Austria
In national parks (IUCN Kat. II) humans theoretically should convert from managers
to observers of natural processes. However, in Europe such parks are mostly small
islands in multiple used landscapes with strong ecological connection to the areas
outside of the park. Wildlife populations, particularly wide ranging red deer, mostly
use habitats that are much larger than the park area. Parks often function as a red-deer
source area for the surrounding region, partly with game-damage problems in
agriculture and forestry. This situation needs a minimum management of red-deer
populations in national parks focused on the aims existing inside as well as outside of
the park. Wildlife management should be oriented on the results of objective
monitoring systems and threshold values which justify management activities in the
park. A park-specific tolerance level for the impact of browsing on forest regeneration
has to be defined because in national parks economic aspects cannot be argued any
more. Operational criteria and indicators are required for monitoring and management.
Ways to manage red-deer populations in Austrian national parks under the conditions
shown above are presented and the following questions are discussed. 1) Can โgame
damageโ be defined for a national park? 2) How can red-deer populations be regulated
by shooting considering the aims of national parks? 3) How is red deer managed in
national parks during winter if natural winter habitats are lacking? 4) Which
experiences exist on the cooperation between wildlife managers of the parks and
wildlife management of neighbouring land owners and hunters?
(Oral presentation.)
175
Wildlife trade in deer species: A need for developing wildlife
forensic techniques.
S. P. Goyal, A. Mandal, R. R. Singh, S. Mishra, and C. P. Sharma
Wildlife Forensic Cell, Wildlife Institute of India, Dehra Dun 248001, India
Efforts for conserving biological resources in South Asia have been one of the major
focus because of richest regions in the world and are the abode of many rare,
endangered and endemic species. Illegal poaching has been one of the major
conservation threats leading to decline of a number species. Of the 830 wildlife
offenses received, 44.8 per cent cases are of deer species. Out of nine deer species in
R 211
India, five species are in Schedule I (Kashmir stag, Thamin, Swamp deer, Mouse deer,
Musk deer) and other four (Chital, Sambar, Hog deer and Barking deer) in Schedule
III of the Wildlife Protection Act, 1972. Most of the items seized in wildlife offences
of deer species are of meat (58%), hair/skin (11.4%), antlers (4.4 %), bone and skull
(3.5) and others. Thus for controlling illegal trade in wildlife parts and products in
Southeast Asia, WII has initiated the work of establishing Wildlife Forensic Facility
for proper implementation of Indian Wildlife (Protection) Act-1972. Hair
characteristics of Indian deer species based on cuticular, and medullar patterns have
been prepared. We have also standardized the protocols of identifying species from
antler based on morphological features, scan electron microscopy and analytical
techniques (TGA, XRD, XRF, ICP-MS). Extraction of DNA is the main challenge for
forensic DNA works. Sometimes the samples are either in very bad quality and
quantity or fixed in formalin which in turn may inhibits Polymerase Chain Reaction.
Isolation of DNA from various tissue samples was done using different protocols viz.
phenol/chloroform (PC) (n=54), DNeasy tissue kit (Qiagen, Germany) (n=100),
Chelex-100 (n=100) and Bio-Robot (n=16) with necessary modifications whenever
required. 28 per cent samples examined (n=275) did not yield any DNA. Quality of
the DNA obtained from these samples was in the order of DNA-Bio robot > DNeasy
tissue kit, (Qiagen, Germany) > PC > Chelex. 48% amplification was achieved in
Polymerase Chain Reaction with mitochondrial cytochrome b universal primers. We
developed the Restriction Fragment Length Polymorphism (RFLP) based profiles for
identifying deer species from meat samples suitable to yield at least 359 bp PCR
products from mt DNA region. Among these restriction enzymes, NlaIII, Ssp1, Stu1
and XhoI are suitable to identify most common Indian deer species. The DNA yield
from some of the degraded meat samples of deer species, antler, and bone was highly
degraded and yield was always < 200 bp. Therefore, we re-designed the PCR primers
and standardized the other protocols for identifying species from such products. We
have also developed DNA based protocols for identifying species from antlers.
(Oral presentation.)
176
Habitat use of pampas deer (Ozotoceros bezoarticus) at agricultural
areas in southern Brazil.
F. G. Braga
Federal University of Paranรก, Rua Saldanha Marinho 1923, Curitiba - 80.730-180,
Brazil
The studied aimed at the evaluation of habitat use by Ozotoceros bezoarticus pampas
deer in modified areas by agricultural activities, in two private estates in Southern
Brazil. The roads of the studied estates were travelled by car through circuits in order
to identify agricultural areas and grasslands, in sixteen monthly field phases, realized
between Feb/01 and May/02. There were computed 1065 observations, the average
212 R
size of the groups was 2.29 + 0.55, and the sexual ratio was 0.83. Thirty-four deaths
were recorded and the main causes of death were predatory actions, hunting and
individuals being run over by motor vehicles. There were recorded high search rates
for oats and wheat (1.0), oats mixed with rye grass (1.0), soy (0.91), and
grasslands(0.87), whereas maize showed the lowest search rate (0.54). Resources
availability was 100% for grasslands, 75% for maize and soy, 43.75% for oats and
wheat, and 31.25% for oats mixed with rye grass. The rates for substratum utilization
were 0.52 for oats, 0.45 for soy, 0.26 for wheat, 0.22 for grasslands, 0.17 for oats
mixed with rye grass and 0.04 for maize. There was spacial segregation between
pampas deer and cattle. Only 14.6% of the sights in the grasslands occurred when
cattle were sharing it. The grasslands proved to be of much importance at the
beginning of the birth peak, whereas soy was intensely used during lactation, and oats
were an important resource during severe frosts. The results of this study indicate that
crops and grazing land are an alternative source of food that may meet certain wants
of the species or act as an alternative source when cattle limits the use of grasslands.
The pampas deer also take into account other features such as rare human interference,
distance to forested areas, also to roads that present heavy traffic and nearness to areas
that may be used as shelter and refuge. The studied population is seriously endangered
and it will not escape extinction unless certain policies to guarantee its conservation
are adopted. These policies should include the handling of areas and control of
pressure vectors.
n unless certain policies to guarantee its conservation are adopted. These policies
should include the handling of areas and control of pressure vectors.
(Poster presentation.)
177
Impact of red deer browsing on the understory of Hungarian
forests.
N. Bleier, K. Katona, L. Szemethy, J. Szรฉkely, M. Nyeste, ร. Fodor, A. Terhes, V.
Kovรกcs, and T. Olajos
St Istvรกn University, Department of Wildlife Biology and Management, St Istvรกn
University, Department of Wildlife Biology and Management, Pรกter Kรกroly u. 1.,
Gรถdรถllล 2103, Hungary
In many European countries large herbivore-forest relationships causes conflicts
between game and forest managers and nature conservers. Overpopulated red deer are
named as primary factor for the forest damages. However, forest habitat improvement
is hardly-considered as a solution against game damages. To describe the problem,
availability of different plant forages and browsing on them by ungulates were
seasonally investigated by sprig counting in the understory of five forested areas in
Hungary. Biomass of available sprigs was also calculated. Our results show that
natural food supply can reach 1500 and 3000 kg per ha during the vegetational period.
R 213
Some species were generally preferred (e.g. locust, elderberry or blackberry), but
many other plant species in the understory was also browsed. Proportion of the
browsed sprigs was always between 0 and 10 percent, but in one area with very low
forage availability this proportion was 35-50 percent. The highest browsing was found
during the vegetational period not in winter. There was no strong relationship between
the proportion of sprigs browsed and the local intensity of area use of ungulates. Our
results show that browsing damages could be much more effectively reduced by
establishing rich understory besides local red deer population control.
(Poster presentation.)
178
Effects of small barriers on habitat use in red deer.
C. B. Sรกnchez-Prieto
1, 2
, J. Carranza
1
, S. Alarcos
1
, and C. Mateos
1
1
Biology and Ethology Unit, Universidad de Extremadura, 10071 Cรกceres, Spain.
2
The Macaulay Instutute. AB15 8QH, Aberdeen, Scotland, UK.
The establishment of fences is a common management practise not only in farming
but also in protected areas. These artificial barriers are mainly aimed to protect
vegetation from the grazing and browsing impact of large mammalian herbivores.
Despite the potential benefit of protecting vegetation and landscape from herbivory,
barriers also constrain animal movements at medium and large scale, which has been
demonstrated to be the cause of serious problems in habitat and species conservation.
However we still know little about their effect on the biology and ecology of plant and
animal communities. The aim of this study is to assess the effect of fences on the
spatial distribution, movement and sociology of Red deer in Doรฑana National Park
(Andalucรญa, South-East of Spain). The study was carried out in an ecotone area
between shrub land and marshland. Red deer daily movements take place between the
shrub land, where they rest, and the ecotone where they forage. Most fences and
human constructions are placed in a way that constrain the movement of red deer
between these two habitats. During the rut of 2003 we carried out censuses in the
ecotone area to assess the effect of adjacent fences . We also tested the effect of
experimentally placed fences. Results from censuses show that areas with natural
access affected by permanent fences or obstacles are less used by red deer. On the
other hand, the experimental placing of barriers produced a decrease in the number of
red deer using the areas of the ecotone affected by the experimental barriers. Also, the
placement of the experimental barriers decreased the number of females per harem on
both areas, affected or non affected by the barriers, showing that any management
during the rut could have unknown and unexpected effects. So, we conclude that the
placing of fences and obstacles in the ecotone cause the underutilization of the
214 R
adjacent habitats. This can produce the overuse of other areas and can affect
parameters of the mating system of the red deer through the effect on the spatial
distribution of females.
(Poster presentation.)
Feeding ecology
216 R
179
Botanical composition of taruka (Hippocamelus antisensis) diet
during rainy season in Huascaran National Park, Peru.
C. Gazzolo
Universidad Nacional Agraria La Molina, Universidad Nacional Agraria La Molina,
Luis Garcia Rojas 175, Urb. Humboldt, Miraflores, Lima 18, Peru
The taruka distributes in the high Andes from north Peru to northwest Argentina. It
lives in mountainous rocky rugged terrain, using also grasslands and scrubland,
between 2500 m in the south to up to 5200 m in the centre and north of its distribution.
Studies on taruka feeding ecology and diet allow us to understand the relationships
between taruka and its ecological niche, as well as the possible overlapping with diets
of free ranging domestic ungulates. The field data were collected on the southwest of
Huascaran NP, central Andes of Peru, throughout puna grasslands and rocky slopes,
during February 2003. The rainy season in the area occurs from December through
March. Samples from all vegetative species were collected, including flowering parts
when present. Tissues were taken from the epidermis of leaves, stems and flowers,
applying them a soft chemical treatment. The tissues were boiled and dyed. The tissue
samples were photographed through the microscope, creating a reference collection.
All fresh pellets found in the field were collected and conserved in alcohol, dried up
in the lab. One gram from each sample was diluted in distilled water and centrifuged,
boiled with NaOH. Sodium hypochlorite was added and the samples were washed on
a 40ยพ mesh. Alcohol was added to the samples and dyed. Fragments were identified
by microscope. Grass species eaten by taruka during the rainy season comprised close
to 57% of the consumed fragments. More than 20 plant species were identified as
eaten by tarukas. Among the species that were represented with more than 4% of the
fragments each, are Poa gymnatha, Bromus villosissimus, Calamagrostis sp., Trisetum
spicatum and Poa spicigera (Poaceae), Luzula racemosa and Distichia muscoides
(Juncaceae), Werneria nubigena and Senecio comosus (Asteraceae), and Ephedra
americana (Ephedraceae, Gymnospermae). This is the first study to find the
importance of grasses for tarukas, which may select them when soft, during the rainy
season. Thus, a possible overlap with domestic ungulates diets should be explored,
eventually helping in the conservation of taruka, and generating an adequate
management of the species and the ecosystem.
(Oral presentation.)
R 217
180
Habitat use by two large deer species (Hippocamelus antisensis and
Odocoileus virginianus) and one small deer species (Mazama
bricenii) in the Apolobamba Integrated Management Natural Area
(La Paz-Bolivia).
A. M. Nuรฑez
BIOTA (Centro de Estudios en Biologรญa Teรณrica y Aplicada), La Paz - Casilla 9641,
Bolivia
The distribution and habitat use of two large deer species, Hippocamelus antisensis
(taruka) and Odocoileus virginianus (luichu), and one small deer species, Mazama
bricenii (chuรฑi taruka), was determined in the Apolobamba Integrated Management
Natural Area (Bolivia). The research was conducted between January and October
2001 and covered an altitudinal range of 2950 to 4760 meters above sea level. In order
to determine the distribution of the three species, direct and indirect observations were
taken, including observation of remains. Study data was also complemented by
surveys carried out with local populations and park guards. Habitat use was
determined via analysis of fecal remains (pellets) collected during transects. Transects
were followed in 10 points, distributed proportionally in four different habitats in
accordance with the area covered by each habitat within the protected area. In each
point, four parallel transects (1km x 2m) were followed, totaling 44 transects. Because
it was not possible to differentiate between H. antisensis and O. virginianus pellets,
all pellets encountered were considered as those of large species. This was not the case
with pellets of M. bricenii as their droppings are easy to differentiate. The four
habitats where the transects were set are: Snowline, Semi-humid High Andean,
Yungas Paramo and Yungas Edge-Cloud Forest. There are no significant differences
in habitat use by the two large deer species. They use three habitats: Snowline, Semi-
humid High Andean, and Yungas Paramo. These results may indicate a sympatric
relationship between the species, particularly in the Semi-humid High Andean habitat,
where climatic and topographic present conditions appropriate for both species. The
small species used only the Yungas Edge-Cloud Forest and was not present above the
highest limits of this habitat.
(Oral presentation.)
218 R
181
Impact of deer browsing and other environmental factors upon
growth and development of fir saplings (Abies alba Mill.) in the
Bieszczady Mountains, southern Poland.
D. Merta and K. Kumรณr
Department of Ecology, Wildlife Research and Ecoturism, Pedagogical University of
Cracov, Podbrzezie 3, 31-054 Krakow, Poland
Growth and development of fir saplings is influenced by deer browsing and by other
factors such as climatic variables, pests, improper planting technique and others.
Therefore, in order to estimate browsing impact of deer population upon fir saplings,
the following experiment in the Bieszczadzy Mountains was carried out. In spring
2000 four fir plantations (1.5 ha each) were established and half of each plantation was
fenced by metal netting. Then using systematic placement 13 sapling plots (2 x 8 m)
in each fenced and each unfenced plantation were established. Each sampling inside
sampling plots was marked with number using plastic sticker. During spring 2002,
autumn 2002 and spring 2003 the following measurements for each sapling were
taken: number of twigs, number of damaged: all twigs, leading shoots and others
twigs, diameter of crown and sapling height.
In spring 2002 sampling plots data showed presence of 1122 saplings (6743/ha) in the
fenced areas and 1062 fir saplings (6 611/ha) in the unfenced ones. This means that
probably due to deer feeding 5.34% of fir saplings died during 2 years after the
plantations were established. From spring 2002 to autumn 2002 mortality of saplings
during growing season was higher in unfenced plots (0.53 % vs. 1.13%).
Measurements in spring 2003 indicated also higher winter mortality of saplings in
unfenced areas (0.90% vs. 1.9%). Spring 2003 survey showed that leading shoot of
0.9% saplings was damaged while in unfenced plots this index was equal to 15.9%.
Saplings from the fenced plots were higher (38.9 cm vs 34.3 cm) and their diameter
of crown radical was larger (0.85 cm vs. 0.75 cm). According to ANOVA test the
above differences were statistically significant. Excluding saplings with the leading
shoot damaged it was documented that significant growth of sapling is reduce if more
than 15% of twigs were damaged. Data analysis between spring 2002 and 2003
provided information, that participation of deer in mortality of saplings is 34.9%, but
remaining 65.1% mortality is caused probably by pest, improper planting technique,
wrong soil preparation and climatic factors. However, deer were responsible for
94.3% damage of leading shoots and for 74.7% of damage of all twigs of saplings. In
the study area population density of red deer amounted to 29.5 animals/1000 ha and
local societies do not agree to reduce number of red deer. Recent publication from the
area showed that chemical protection of sapling is much less effective than fencing.
Therefore in order to mitigate conflict between deer management and forest
management fencing of small size young plantation of fir would be most suitable in
the Bieszczady Mountains.
(Oral presentation)
R 219
182
Why deer strip bark? -two case studies of bark stripping by sika
deer in central Japan.
M. Ando
1
, Z. Jiang
2
, and E. Shibata
3
1
Kyoto Prefectural Forestry Experimental Station, Forest Management Section,
Kameoka city, Kyoto pref., 621-0851, Japan
2
Wildlife Management Office Inc., Japan
3
Forest Protection Lab., Graduate School of Bioagricultural Sciences, Nagoya Univ.,
Japan
Bark stripping by cervids occurs worldwide. In many cases, the occurrences of bark
stripping are frequent in winter and early spring, and explained that are caused by food
shortages. On the other hand, in some cases, cervids seems to eat bark without food
shortages. In Japan, sika deer, Cervus nippon, also strip bark. We give a presentation
about two investigations of the bark stripping by sika deer in central Japan. We carried
out our studies held at two regions to clarify the relationships between the seasonal
changes of bark stripping and food quality. These investigations gave contorastive
results. (1) In Mt. Ohdaigahara, bark stripping is most intensive during summer (July
to September), when Sasa nipponica, the deer's main forage in here, is abundant. It
suggests that bark stripping is not due to food shortages. The nutritive value of bark
is lower with much lignin, and that of S. nipponica, is higher with much crude protein
and hemicellulose contents in summer. Sika deer seems to eat the bark to balance the
digestible nutrients of summer forage. (2) In Northern Mount Fuji district, planted
veitch fir were stripped the bark. Bark stripping occurred from December to May or
June with a peak in March April. The stripping period overlapped with the periods of
low food availability and poor food nutrition of sika deer, from January to April.
February is the worst forage period in terms of both quantity and quality. There was
a time lag of 1 to 2 months in the peak of bark stripping in March April when
compared with the poor forage period in February. This time lag may suggest that sika
deer need more nutritious and easily digested food from March due to increased
nutrient demands that result from depleted body condition in both sexes, gestation of
pregnant females, and the recovery of active metabolism. These two cases suggest that
there are several seasonality and reasons of bark stripping by sika deer in each habitat
although they are the same species, and also suggest that the quality and quantity of
other available foods are concerned with the bark stripping occurrence.
(Poster presentation.)
220 R
183
Influence of an extreme climatic event on the winter diet of red and
roe deer in northeastern France.
D. B. F. Storms, S. Said, J.-L. Hamann, C. Saint-Andrieux, J.-L. Wilhelm, and F.
Klein
Office National de la Chasse et de la Faune Sauvage, Centre National d'Etudes et de
Recherche Appliquรฉe Cervidรฉs-Sanglier, 1 Place Exelmans, 55000 Bar le Duc, France
Extreme climatic events may profoundly affect herbivore population dynamics and
habitat use by returning tracts of mature forests to early seral stages where food and
cover resources rapidly develop. In December 1999, hurricane Lothar struck France,
Switzerland and Germany, and caused widespread destruction to forests. Studies
carried out in France suggest that roe deer (Capreolus capreolus) may have benefited
from the creation of hurricane-related clearings, at least in the short term. It has been
suggested that the combination of a reduction in hunting due to huntersโ inability to
access the areas and an improvement in habitat characteristics in clearings resulted in
Lothar having a positive effect on roe deer population dynamics. Here, we studied the
winter diet of red (Cervus elaphus) and roe deer before and after hurricane Lothar in
a heavily damaged forest, assuming that the hurricane, by creating numerous clearings
in which food availability for herbivores increased rapidly, allowed deer to specialize
on preferred plant species that benefited from habitat modifications, particularly
bramble and grasses. According to their feeding types and ecology, we expected that
the increase in the availability of grasses and bramble after the hurricane would result
in an increase in the proportions of these plants in the diet of red deer and an increase
in the proportion of bramble in the diet of roe deer. Analysis of stomach contents
revealed that consumption of grasses by red deer almost doubled following the
hurricane, but no effect on bramble consumption by either deer species could be
detected. Our study also revealed a significant annual variation in bramble
consumption, which suggests that its availability as a food resource for deer varies
from year to year, and this may have masked the hurricane effect. We then discuss the
role of disturbances in improving habitat for forest ungulates, and the role of key
alternative food resource in helping forest managers to find solutions for mitigating
ungulate impact on tree regeneration.
(Poster presentation.)
Seasonal and non-seasonal
deer: (Arctic to Tropic)
222 R
184
Seasonal migration pattern of red deer ( Cervus elaphus L.) in the
central Slovakian mountains.
S. Finฤo
1
, J. Buฤko
1
, and S. Steyaert
2
1
National Forest Centre, T.G.Masaryka 22, 96092 Slovakia
2
Wageningen University,Droevendaalsesteeg 51,6708 PB-Wageningen, Holland
Red deer is a common species and the current contigous range encompasses 39,054
km
2
, or 80 % of the country with numbers estimated up to 38,000 individuals. A
proportion of red deer in the western Carpathian mountains seasonally migrate
between wintering areas and higher altitude grazing grounds. In 1998 2005, the
movement and migration pattern of 13 red deer in the Nรญzke Tatry and Poana
mountains were studied by VHF telemetry. The study area elevations ranged from 430
metres to 1,660 metres and composed of a great diversity of forest ecosystems. In total
8 stags and 5 hinds were individually
radio-tracked, the duration of telemetry varying from 6 to 56 months. Spatiotemporal
behaviour of red deer was described by 6 characteristics: migratory status, migration
date, size of seasonal home range, average distance between seasonal ranges, altitude
of telemetry locations and direction of migration. Migration was defined as the date
upon which migrant red deer left their seasonal home range to the date they
established their summer home range. 5 stags migrating from winter range for non-
overlapping summer ranges had migration dates from April 11 to May 4 and return
dates in the fall ranged from November 11 to January 17, which particularly depended
upon the depth of snow covering the summer ranges. Spring migration peaked and
concentrated more than the fall migration. Two stags and five hinds were classified
as resident and a young stag classed as a transitional type, between migrant and
resident, used a winter home range which overlaped an extrordinary large summer
home range. The size of the seasonal home ranges was derived by an MCP estimator.
Migrants had larger winter and summer ranges than residents (winter 13.2 km
2
vs.
5.6 km
2
, summer 11.2 km
2
vs. 3.2 km
2
) with the size of winter ranges differing
significantly. Surprisingly the size of winter and summer home ranges (migrants
13.2 km
2
vs. 11.2 km
2
; residents 5.6 km
2
vs. 3.2 km
2
) were not statistically different.
Daily movements in the search for widespread supplementary food, possibly increased
and enlarged winter home ranges. Average distances between the centres of summer
and winter ranges were significantly larger for migrants (9.4 km; range 2.9-14.7 km)
than for residents (0.9 km; range 0.1-2.6 km). It appears that spring migration can be
well characterised by both change of altitude and direction of movement. It was
obvious that in spring red deer migrated from lower winter ranges to higher altitudes
and that their direction of movement was also important as indicated by the span of
bearings which ranged from 308 to 100 approximately northwest to east.
(Oral presentation.)
R 223
185
Scale-dependent habitat selection of GPS-collared Alpine red deer
the role of food availability and quality.
B. Zweifel-Schielly and W. Suter
Swiss Federal Research Institute WSL, Zuercherstrasse 111, CH-8903 Birmensdorf,
Switzerland
Understanding habitat selection usually requires a scale-conscious approach. On the
behavioural level, habitat selection may be seen as an animals trade-off between
responding to different needs such us feeding, social contacts, predator avoidance, and
others. Habitat selection in mountain ungulates may be especially fine-tuned as
difficult topography and harsh climate often restrict access to food resources with
sufficient nutritional value. We studied variation in habitat and diet selection of GPS-
collared red deer Cervus elaphus in the northern Swiss Alps, and measured both
availability and use of food resources with respect to biomass and nutritional value
over an area of 250 km
2
. We defined selectivity at two scales, home-range scale and
site scale. We hypothesized that during winter, selectivity would be strongest, and that
requirements for highly nutritious food would be the main driving factor in habitat
selectivity across seasons and spatial scales. At both scales, selectivity in habitat
selection decreased from winter to summer. Forest was consistently preferred over
open land at home-range scale, while at site scale, preference for open land declined
from winter to summer. With respect to forest structures, red deer exhibited seasonally
different preferences for structural properties in choosing home ranges. About half of
all preferences were associated with habitat offering better forage, either in terms of
higher biomass or higher content of crude proteins, or both. Overall diet was
dominated by graminoids (41%), but tree browse (29%) and forbs and Rubus (18%)
were also important. Variation in diet composition was mostly associated with season,
but effects of habitat at home-range scale and sex were also detected. Diet selection
was consistent with the findings from the analysis of habitat selection, reflecting
strong use of plant groups with high biomass, easy availability and high nutritional
quality (low cellulose/lignin, low lignin, high organic matter, and high crude protein
contents).
(Oral presentation.)
224 R
186
Photic modulation of the temporal pattern and rate of activity in
reindeer.
B. E. H. van Oort
1
, N. J. C. Tyler
2
, M. P. Gerkema
3
, L. Folkow
1
, and K. A. Stokkan
1
1
Department of Arctic Biology, University of Tromsรธ, Norway
2
Centre for Sami Studies, University of Tromsรธ, Norway
3
Department of Chronobiology, University of Groningen, The Netherlands
Northern species of deer have adapted to the relatively predictable seasonal variation
in food supply characteristic of temperate environments through the evolution of
annual cycles of appetite and growth. These cycles persist under constant conditions,
indicating their endogenous basis, but are normally entrained by the annual cycle in
day length. Several species also display a pronounced annual cycle in the daily rate
of activity (h.24h-1). Increased activity in summer reflects the permissive effect of a
long photo period on activity and, in particular, on time spent feeding while marked
peaks of activity around dawn and dusk indicate that the activity pattern is under photo
periodic control. Reindeer/caribou live at high latitudes where the light/dark (LD)
cycle of night and day is weak or absent for much of the year and several reports
suggest that neither the temporal pattern nor the rate of activity in this species is under
photoperiodic control. These results, if confirmed, would indicate that the activity of
reindeer may be independent of the seasonal fluctuations in metabolic demand
associated with the annual cycle of growth. We examined activity in reindeer in
relation to ambient photic conditions using continuous records collected across one
calendar year in free-living Rangifer tarandus tarandus (NR) and R. t. platyrhynchus
(SR) at 70
o
and 78
o
N, respectively. In both sub-species, alternating ultradian bouts of
activity and inactivity persisted across the 24h day throughout the year. However, in
contrast with earlier suggestions, the LD cycle, when present, imposed a diel rhythm
in the otherwise continuous 24h pattern. All the reindeer displayed a pronounced
annual cycle in the daily rate of activity which was closely in phase with the appetite
cycle. The amplitude (A) of the activity cycle was greater in SR (A = 2.4h) compared
with NR (A = 1.5h) reflecting the larger amplitude of the appetite cycle in this sub-
species. We conclude that the monthly rate and the 24h pattern of activity in reindeer
are modulated by the photic environment. Weaker diel rhythmicity in SR compared
with NR results from a latitudinal decrease in circadian organisation and photic
responsiveness in Rangifer.
(Oral presentation.)
R 225
187
Habitat use and selection of fallow deer (Dama dama L.) in a
Mediterranean environment.
P. Di Luzio, P. Montanaro, and S. Focardi
Istituto Nazionale per la Fauna Selvatica, Via Cร Fornacetta,9 40064 Ozzano dell
Emilia (BO), Italy
The distribution and quality of resources as well as the habitat structure play an
important role in determining the spatial behaviour of deer. The knowledge of the
relationship between habitat and ranging behaviour is essential for the management
of wild populations. At the present time the ecological requirements and habitat
selection of fallow deer (Dama dama L.), in spite of its wide distribution, is scarcely
known especially in Mediterranean region where is the original range of the species.
We studied the habitat use and selection of fallow deer in the Castelporziano Estate
near Rome (Italy) characterized by a Mediterranean climate with mild winters and
warm and dry summers. The vegetation is mainly composed by evergreen oak forest
(44%), deciduous and mixed oak forest (21%), pine woods (7 %), Mediterranean
maquis (20%) and open areas (8%). Twenty-two (13 males and 9 females) adult
fallow deer were caught, from January 2000 to December 2003, in vertical nets, in
cage traps or immobilized by using a dart gun and fitted with radio-collars. We
collected 6638 locations in 4 years, and those with a 95% confidential ellipse greater
than 1 ha was discharged (33,5%). Compositional analysis was performed in order to
assess the habitat use and selection a two kind of geographical level: 1) home range
vs. study area; 2) location vs. home range. Data were investigated at two different
temporal scales (annual and seasonal). Differences between sex, daytime period, and
activity state were tested. At annual scale only differences between sexes emerged. At
II level of selection females avoided open areas whereas males evaded pine woods.
Therefore, at seasonal scale, both differences among sexes and year resulted. Females
selected maquis and evergreen oak forest, while males preferred deciduous oak forest
and open areas. At the II level of selection males actively preferred open areas that
were placed at the lowest rank by the females.
(Poster presentation.)
226 R
188
Function of habitat segregation in regulation of isolated sika deer
population.
S. Tatsuzawa
Hokkaido University, Hokkaido University, Department of Regional Sciences,
Graduate School of Letters, Japan
In ungulate populations, there are many reports on the phenomenon of habitat
segregation, but little on its function in population dynamics. To study the mechanism
and function of habitat segregation in population dynamics, a long-term seasonal
census has been carried out for fifteen years on a Mage sika population Cervus nippon
mageshimae of Mage-shima island (31N, 131E, 12 square-km), which has been
maintained at least for a thousand years without predators, competitors, snowfall, nor
human disturbances. In this closed and long-maintained population, obvious habitat
segregation in sex-age classes was observed ordinarily, namely females and fawns use
forest area, prime males open-rich grassland, and yearling males open-poor grassland
at seashore. The total density (13-52 animals/square-km) and its seasonal variation
(SD=.16-.29) have fluctuated yearly, and a density-depended regulation has detected
in this population. This strict and agile density fluctuation has brought by drastic
changes in natality (10.9-62.9 fawns/females) and mortality of yearling males (0-36.1
%), and these two demographic parameters were skewed spatially corresponding to
the property of habitat segregation in sex-age classes. This tendency of spatial skew
in the mortality was remarkable especially in โsinkedโ years. Consequently, spatial
segregation in sex-age classes is considered to reinforce the density dependence in this
closed population, and hence, to contribute higher sustainability to this small insular
deer population.
(Poster presentation.)
Venison and its potential
contribution to diet
228 R
189
Venison and the history of early European hunting enclosures.
T. J. Fletcher
Reediehill Deer Farm, Auchtermuchty, Scotland
The prey of hunting primates is often supposed to have been pivotal in the evolution
of large brained hominids. This was not only as nutrition but also because, as perhaps
the first valued possession, meat endowed the successful hunter with the power of
bestowing gifts and bribes. Thus the prestige associated with a productive hunt has a
long ancestry. In this context the prodigious effort made by rulers throughout Eurasia
from prehistory to modern times to ensure prolific hunts becomes comprehensible.
One of the best ways to achieve this was by the creation of the hunting reserve or
enclosed deer park. Recent hypotheses on the parkland nature of the original European
woodland suggest that โhagaโ were barriers surrounding groves. In Anglo-Saxon
England the hayโ was a deer enclosure. By the time of the 11th century Norman
invasion there were several hundred such hays which became known by the Latin
word โparkโ which in itself shares roots with the Old Persian word โpairidaezaโ. The
Persian paradises not only became associated with Eden, heaven and a state of
supreme bliss but also, essentially, incorporated hunting preserves.England, and to a
lesser extent Scotland, would appear to be unique in the number of deer parks which
were created during the medieval era, two to three thousand at a time when the human
population was only around three to four million. England still possesses more deer
parks than any other region in the world. Since all deer belonged to the monarch the
establishment of a deer park depended on royal patronage. Venison from those parks
could not be sold until the eighteenth century but was frequently gifted, echoing the
โgiftsโ of prey made by the hunting primates.Many parks existed for almost a
thousand years evolving to meet contemporary fashions and becoming the designed
landscapes which have been described as Englandโs greatest contribution to art. This
paper discusses the origin and development of the British deer park, the ways in which
they were used and their deer managed to produce venison, and their present and
future value.
(Oral presentation.)
R 229
190
Fatty acid profiles in Javan rusa (Cervus timorensis russa) stags.
R. Sookhareea
1
, R. Tume
2
, W. R. Shorthose
2
, and G. M. Dryden
3
1
Ministry of Agriculture, Food Technology and Natural Resources, Reduit, Mauritius
2
Food Science Australia, Brisbane
3
School of Animal Studies, University of Queensland, Gatton, Queensland 4343,
Australia
Rusa deer are an important commercial species in the subtropics and tropics
(Sinclair, 1997; Maudet, 1999). However, although they are reared almost exclusively
for venison, there are few data on the quality of their meat. The fatty acid profile of
meat influences its palatability, proneness to rancidity and nutritional value. There are
several reports of the fatty acid profiles of other deer species (e.g. Manley and Forss,
1979; Rule and McCormick, 1998; Ishida et al., 2001; Volpelli et al., 2003) but none
for rusa deer. This experiment was conducted to investigate the fatty acid profiles of
the intramuscular lipid and fat depots of rusa venison. Because castration will control
undesirable behaviour in stags and may improve dressing percentage without affecting
growth in young stags (Sookhareea et al., 2001) we also investigated the effects of
castration on venison fatty acids.
Rusa stag calves, born in autumn (March), were surgically castrated after weaning
(four animals) or left entire (seven animals) and the effects of these treatments on fatty
acid profiles were examined in deer slaughtered at 25 months of age. The deer grazed
pastures of Pennisetum clandestinum and Trifolium repens from autumn to early
spring, and of predominantly Cynodon dactylon and Panicum maximum in late spring
and summer. Supplements of Medicago sativa and grass hay and sorghum (Sorghum
bicolor) grain were given during periods of pasture shortage in the winter. Details of
the animals and management are given in Sookhareea et al. (2001b). Deer were
slaughtered and processed using standard procedures (Sookhareea et al., 2001a).
Samples of visceral, kidney, subcutaneous, and intermuscular fat were taken after
processing the carcase. At 24 h postmortem, the m. longissimus dorsi (LD) was
excised and samples taken for fatty acid analysis. All samples were stored frozen (-20
o
C) until analysed. Fatty acids were determined by GLC procedures. Total lipids from
the sample tissue were extracted from the homogenates of 0.5 to 1g samples using
chloroform-methanol (2:1 v/v) according to Folch et al. (1957). The lipid extracts were
methylated (5 % sulphuric acid in methanol) overnight at 60
o
C and the FAME were
extracted with petroleum spirit, evaporated under nitrogen and diluted. FAME were
isolated by gas-liquid chromatography (Packard-Becker Model 419 fitted with a flame
ionisation detector and a 50 m ร 0.25 mm Chrompack CP-Sil-88 capillary column, and
using N
2
as the carrier gas), using standard FAME mixtures (Sigma, St. Louis, MO)
as standards. Data were subjected to analysis of variance using the general linear
model (Statistical Analysis Systems, 1989).
Fatty acid profiles are presented in Tab. 1. Castration had no effect on any fatty
acid profile. The depot fats were significantly (P<0.01) more saturated than the
intramuscular lipid. Palmitate plus stearate was 62% in kidney fat and 43 to 46% in
230 R
the other depot fats, compared to 34% in the intramuscular lipid. In all depots and in
the LD intramuscular lipid the quantitatively important fatty acids were palmitic (23
to 34% of detected fatty acids), stearic (12 to 28%) and oleic (14 to 25%) acids.
Intermuscular and subcutaneous fats had most (24%) oleic acid, significantly (P<0.01)
more than the other depot fats (15 and 19%) and the intramuscular lipid (14%).
Saturated fatty acids (SFA) and monounsaturated fatty acids (MUFA) were
respectively 40 to 70% and 26 to 46% of the total fatty acids. Polyunsaturated fatty
acids (PUFA) were approximately 2% of the depot fatty acids but were 23% of the
intramuscular lipid. Traces of trans fatty acids (16:1t and 18:1t(11)) were found in all
of the fat depots and the intramuscular lipid.
A saturation gradient was apparent in the depot fats, with the fatty acids in the
interior depots (kidney and visceral fats) being more saturated than those in the
intermuscular and subcutaneous fats. Much (72.5 and 67.1% respectively) of the
kidney and visceral fats was saturated and there were only traces of PUFA. The
subcutaneous fat was the least saturated depot fat; SFA were 51.1% of the total fatty
acids.
In contrast to the depot fats, the intramuscular lipid of LD had significantly
(P<0.01) more unsaturated fatty acids. The MUFA contained palmitoleic (12.0%) and
oleic acids (13.9%). While linoleic and linolenic acids were found in the other tissues
in trace amounts, they occurred in the intramuscular lipid at 11.3 and 3.3%. In contrast
to the depot fats, the long-chain polyunsaturated acids arachidonic (6.3%) and
docosapentaenoic (1.8%) acids were detected in the LD.
This study is consistent with data from red deer and reindeer (Garton and Duncan,
1971) and white tailed deer (Rule and McCormick, 1998) in that it showed that the fat
depots of deer are highly saturated. The saturation gradient observed in our experiment
has been reported previously in steers (Ashes et al., 1993) and deer (Rule and
McCormick, 1998).
The fatty acid composition of the intramuscular lipid in the LD of rusa stags was
similar to other deer species (Manley and Forss, 1979; Sinclair et al., 1982; Ishida et
al., 1991; Volpelli et al., 2003). It is interesting that the fatty acid profile for these rusa
deer resembled that of sambar (C. unicolor) deer in the study of Sinclair et al (1982).
Rusa and sambar are genetically close (Emerson and Tate, 1993) and they may have
similar capacities for chain elongation and desaturation.
The higher proportion of PUFA in the intramuscular lipid than in the depot fats can
be attributed to the relative contributions of triglycerides (high SFA, low PUFA) and
membrane phospholipids (low SFA, high PUFA) in these tissues (Manley and Forss
1979). The presence of appreciable quantities of PUFA in the LD suggested that some
dietary PUFA must escape microbial hydrogenation, as the linoleic and linolenic acids
must have originated from the diet. The presence of 18:1 isomers reflects the synthesis
of these by rumen bacteria (AbuGhazaleh et al., 2005).
Other studies have shown that the intramuscular lipid of venison contains more
PUFA than other livestock species (Sinclair et al., 1982; Ishida et al., 2001; Rule et
al., 2002). The main SFA in the intramuscular lipid were palmitic and stearic acids,
with some myristic acid. Palmitic and myristic acids are considered undesirable as
they increase LDL cholesterol in humans if the diet is low in linoleic acid (Grundy,
1994; French et al., 2002). On the other hand, this muscle has high proportions of
MUFA and PUFA, which are nutritionally desirable. The fatty acids of the fat depots
R 231
are highly saturated, but as these are primarily extra-muscular and can be removed by
trimming, this feature of venison need not reduce its overall desirability.
Table 1. Fatty acid composition (least squares means, % of total fatty acid) of the
intramuscular lipid of m. longissimus dorsi, and the kidney, visceral, intermuscular
and subcutaneous fats of rusa stags, reared on pasture and slaughtered at 25 months.
Fatty
acid
Fat depots Longissi-
mus dorsi sem
Kid-
ney Visceral Inter-
muscular
Sub-
cutaneous
Saturated fatty acids
12:0 0.4a 0.3b 0.3b 0.2b -- 0.02
14:0 7.7a 8.5a 7.5a 6.4b 4.3c 0.34
15:0 1.3a 1.1b 1.1b 1.0b 0.4c 0.08
16:0 33.6
a
32.9a 29.0b 29.4b 22.9c 0.44
17:0 1.5a 1.1b 1.0b 0.8b 0.4c 0.09
18:0 28.0
a
23.2b 15.4c 13.3d 11.5e 0.90
Monounsaturated fatty acids
14:1 0.4c 0.9c 2.1b 1.7b 2.9a 0.25
16:1t 0.6a 0.4b 0.6a 0.6a 0.4b 0.05
16:1c 2.7b 4.5b 9.5a 10.2a 12.0a 0.57
18:1c(9) 14.5
c
18.5b 23.2a 25.1a 13.9c 0.69
18:1t(11) 6.0a 4.0b 3.4b 3.5b 1.1c 0.24
18:1c(11) 1.5d 2.9d 4.3c 5.2b 6.9a 0.24
Polyunsaturated fatty acids
18:2c 0.9b 1.1b 1.2b 1.4b 11.3a 0.33
18:3c 0.9b 0.8b 0.9b 0.9b 3.3a 0.11
20:4 -- -- -- -- 6.3
232 R
Fatty
acid
Fat depots Longissi-
mus dorsi sem
Kid-
ney Visceral Inter-
muscular
Sub-
cutaneous
22:5 -- -- -- -- 1.8
We thank Prof. K.A. Woodford for making these deer available, and he and Dr.
D.G. Taylor for helpful advice. Mr. R. McAllister assisted with slaughtering and
carcase processing, and we thank Mr. T. Larsen for assistance with the fatty acid
analyses.
References
AbuGhazaleh, A. A., Riley, M. B., Thies, E. E., Jenkins, T. C. (2005) Dilution rate
and pH effects on the conversion of oleic acid to trans C18:1 positional isomers in
continuous culture. Journal of Dairy Science 88:4334-4341.
Ashes, J. R., Thompson, R. H, Gulati, S.K., Brown, G. H., Scott, T.W., Rich, A. C.,
Rich, J. C. (1993) A comparison of fatty acid profiles and carcass characteristics of
feedlot steers fed canola seed and sunflower seed meal supplements protected from
metabolism in the rumen. Australian Journal of Agricultural Research 44:1103-1112.
Emerson, B. C., Tate, M. L. (1993) Genetic analysis of evolutionary relationships
among deer (subfamily Cervinae). Journal of Heredity 84:266-273.
Folch, J., Lees, M., Stanley, G. H. S. (1957) A simple method for the isolation and
purification of total lipids from animal tissues. Journal of Biological Chemistry
226:497-509.
French, M. A., Sundram, K., Clandinin, M. T. (2002) Cholesterolaemic effect of
palmitic acid in relation to other dietary fatty acids. Asia Pacific Journal of Clinical
Nutrition 11:S401-S407.
Garton, G. A., Duncan, W. R. H. (1971) Fatty acid composition and intramolecular
structure of triglycerides from adipose tissue of the red deer and the reindeer. Journal
of the Science of Food and Agriculture 22:29-33.
Grundy, S. M. (1994) Influence of stearic acid on cholesterol metabolism relative to
other long-chain fatty acids. American Journal of Clinical Nutrition 60:986S-990S.
Ishida, M., Ohno, H., Takeda, T., Ikeda, S., Saito, T. (1991) General composition and
depot fats characteristics Japanese Sika Deer (Cervus nippon) carcass. Animal Science
and Technology 62:904โ908.
Ishida, M., Odashima, E., Ikeda, S., Takeda, T. (2001) Comparison of Japanese deer
meat and cattle meat for cholesterol level and fatty acid composition. Nippon
Shokuhin Kogaku Kaishi 48:20โ26.
R 233
Manley, T. R., Forss, D. A. (1979) Fatty-acids of meat lipids from young red deer
(Cervus elaphus). Journal of the Science of Food and Agriculture 30:927-931.
Maudet, F. (1999) Evolution, systรฉmatique et rรฉpartition du cerf rusa. In S. Le Bel, F.
Maudet, N. Barrรฉ, D. Bourzat (eds.) Le Cerf Rusa en Nouvelle Caledonie, Actes dโun
seminaire, Port Laguerre. pp. 4-8.
Rule, D. C., Broughton, K. S., Shellito, S. M., Maiorano, G. (2002) Comparison of
muscle fatty acid profiles and cholesterol concentrations of bison, beef cattle, elk, and
chicken. Journal of Animal Science 80:1202-1211.
Rule, D. C., McCormick, R. J. (1998) Fatty acid composition and cholesterol
concentration in tissues of white-tailed deer (Odocoileus virginianus) as influenced
by lactation, age, and season of the year. Comparative Biochemistry and Physiology
Part B: Biochemistry and Molecular Biology 119:563-570.
Sinclair, A. J., Slattery, W. J., OโDea, K. (1982) The analysis of polyunsaturated fatty
acids in meat by capillary gas-liquid chromatography. Journal of the Science of Food
and Agriculture 33:771-776.
Sinclair, S.E. (1997) Deer farming in Queensland โ 1997 farm survey report.
Queensland Department of Primary Industries, Brisbane.
Sookhareea, R., Taylor, D.G., Dryden, G. McL., Woodford, K. A. (2001a) Primal
joints and hind-leg cuts of entire and castrated Javan rusa (Cervus timorensis rusa)
stags. Meat Science 58:9-15.
Sookhareea, R., Woodford, K. A., Dryden, G. McL. (2001b) The effect of castration
on growth and body composition of Javan rusa stags. Asian-Australasian Journal of
Animal Sciences 14:608-614.
Statistical Analysis Systems (1988) SAS/STATยฉ User's Guide Release 6.03 Edition.
SAS Institute Inc., Cary, NC.
Volpelli, L. A., Valusso, R., Morgante, M., Pittia, P., Piasentier, E. (2003) Meat
quality in male fallow deer (Dama dama): effects of age and supplementary feeding.
Meat Science 65:555-562.
(Oral presentation.)
234 R
191
The effect of pelvic suspension on the biochemical and sesnory
quality of venison from red deer (Cervus elaphus) and fallow deer
(Dama dama).
C. L. Hutchison, J. S. Flesch, and R. C. Mulley
University of Western Sydney, Penrith South DC NSW 1797, Australia
The need to link carcass production with eating quality has long-term implications
for acceptance of venison as a favoured consumer selection. Hence, definition of the
link between post slaughter carcass management and the relationship with cooking and
eating quality will increase opportunities for target marketing, which should increase
farm profitability and consumer satisfaction if product consistency is enhanced. It is
acknowledged that factors such as body condition, pre slaughter management,
methods of slaughter and methods of meat storage can have a significant impact on
eating quality of the final product (Mulley et al, 2006). Texture, flavour and
tenderness are attributes valued by consumers as very important in relation to the
eating quality of meat (Lundesjo et al 2003). Different populations of consumers have
different preferences for these quality attributes, something that affects the market for
all types of meat. However, regardless of the consumer group, the consistency of meat
quality is very important, and the product should be of the same quality every time it
is purchased. In the Australian beef grading system Meat Standards Australia (MSA)
these consumer important sensory quality attributes have been weighted in an overall
score where tenderness represents 40%, flavour 20%, juiciness 10% and overall liking
30% (MSA, 2001).
Tenderness is an important meat quality characteristic for consumers. Muscles in
the butt and loin of a carcass can be restrained from shortening (toughening) by
hanging the whole carcass by the aitchbone (obturator foramen) or the pelvic ligament.
This process is referred to commercially as tenderstetching or pelvic suspension, and
has been shown to increase meat tenderness in several other species (Sims et al 2004).
Tenderness is one of the most important parameters as rated by consumers, in terms
of eating quality. Major factors which affect tenderness include cut of meat, animal
age, cold shortening that can occur during chilling and pre-slaughter animal stress
leading to high pH. Toughness can be prevented by the use of techniques such as
electrical stimulation, which accelerates rigor and pH decline (Wiklund et al 2001),
or hanging the carcass in such a way that muscles will be stretched and not allowed
to contract, hence the term 'tenderstretch'. Pelvic suspension, if found to be beneficial
in producing consistently tender venison may be a useful alternative technique to
electrical stimulation in Australia, where electrical stimulation is generally unavailable
for deer processing.
In this study post slaughter management of carcasses compared Achilles tendon
and pelvic suspension methods of hanging to determine effects on venison quality.
Red stags (n=14) and fallow deer bucks (n=20) and fallow deer does (n=10) of
between 12 and 36 months of age and in prime slaughter condition were examined.
R 235
Animals were assessed ante-mortem to determine suitability of coition for slaughter.
They were then slaughtered by captive bolt stunning and exsanguinated within 3
seconds of the stun. Carcasses were split down the spine and assigned to one of the
two hanging methods. Carcasses were then evaluated for pH (decline and ultimate)
and core body temperature. After 24 hours samples were removed and vacuum
packaged prior to being frozen at -21ยฐC. Once thawed, samples were analysed in
triplicate for intra muscular fat (Soxhlet method), tenderness (Warner Bratzler Shear
force), colour (Minolta Lab), moisture (air oven) and freeze/thaw stability.
There are a number of biochemical measurements commonly used by meat
scientists for meat description that can also indicate changes to meat quality for eating,
cooking, storage and processing. It is well known for instance that muscle pH is
associated with muscle tenderness. The colour of meat is an important characteristic
for marketing, as customer selection is often associated with the appearance of the
product. Water holding capacity is another characteristic of meat that is connected to
consumer perception.
Descriptive and consumer preference sensory testing was undertaken with 42
panelists. There was a balance of male and female participants and a balanced age
distribution from the target market, being 25 to 55 years of age. Panellists were
screened to determine if they were eaters of red meat and were willing to try venison
or were current venison consumers. Panellists were asked to refrain from smoking one
hour prior to and during the sessions. Panellists undertook a familiarisation session to
assist in identifying quality parameters for venison i.e. liver/game flavour, colour,
tenderness and juiciness and use of the survey tool. Samples were prepared to reach
an internal temperature of between 68-72ยฐC, which is determined to produce a product
which remains palatable and safe for consumption. Panellists were presented the
sample identified by random three digit codes and answered questions on the
descriptive test by indicating on a 10cm line scale how they rated the sample for
flavour, colour, juiciness, tenderness and overall liking. Panellists were seated in
individual isolation booths with a drinking cup of water (90%) and apple juice (10%)
to cleanse the palate between tastes.
There were no statistical differences between Achilles hung and tenderstretched
carcasses for the meat quality parameters of pH, colour, moisture or fat. While there
was no detectable difference of hanging method on raw shear (p=0.06),
tenderstretched carcases had significantly lower cooked shear force values than
Achilles hung carcasses (p =<0.001). (Table 1)
Table 1: Meat quality attributes of fallow bucks hung by the Achilles tendon and
Pelvic Suspension method for M. Longissimus dorsi.
Hanging Achilles Hung Pelvic Suspension
PH 5.80a (0.04) 5.80a (0.03)
Cooked Shear(g) 5889.5a (341.7) 4402.2b (157.8)
Raw Shear(g) 2177.4a (115.2) 2598.4a (165.6)
236 R
Hanging Achilles Hung Pelvic Suspension
Colour L* 20.46a (0.39) 21.21a (0.55)
Colour a* 12.29a (0.52) 11.56a (0.36)
Colour b* 0.088a (0.20) 0.117a (0.11)
Moist (%) 75.70a (0.17) 76.02a (0.15)
Fat(%) 2.74a (0.16) 3.06a (0.24)
Water (%) 13.56a (0.98) 13.69b (0.76)
Means and standard error of means (in parenthesis) are shown Treatments followed
by the same letter in the rows are not significantly different (p<0.05)
Fallow does exhibited a significant difference between methods of suspension for
cooked shear (p=0.015) but not for other parameters tested. (Table 2)
Table 2: Meat quality attributes of fallow does hung by the Achilles tendon and Pelvic
Suspension method for M longissimus dorsi.
Hanging Achilles Hung Pelvic Suspension
pH 5.73a (0.03) 5.69a (0.02)
Cooked Shear (g) 4558.6a (217.2) 3778.6b (155.9)
Raw Shear (g) 2545.9a (277.3) 2721.6a (271.5)
Colour L* 22.13a (0.45) 22.24a (0.43)
Colour a* 13.56a (0.39) 13.56a (0.37)
Colour b* 1.68a (0.28) 1.68a (0.35)
Moist % 75.49a (0.47) 74.35a (0.28)
IM Fat % 1.78a (0.18) 1.89a (0.17)
Freeze Thaw loss % 16.67a (1.04) 13.76a (1.08)
Treatments followed by the same letter in the rows are not significantly different
(p<0.05)
In the red stags there was a significant difference between carcasses hung by the
Achilles tendon compared with pelvic suspension for cooked shear (P<0.001).
(Table 3)
R 237
Table 3: Meat quality attributes of carcasses of red deer stags hung by the Achilles
tendon and Tenderstretch method after slaughter for M. Longissimus dorsi.
Hanging Achilles Hung Pelvic Suspension
PH 5.63a (2.67) 5.63a (2.67)
Cooked Shear (g) 5475.8a (298.1) 4124.1b (154.5)
Raw Shear (g) 3535.1a (184.0) 3761.8a (167.9)
Colour L* 23.01a (0.39) 23.19a (0.28)
Colour a* 2.96a (0.22) 3.05a (0.15)
Colour b* 2.96a (0.22) 3.05a (0.15)
Moist % 75.95a (0.14) 75.96a (0.2)
IM Fat % 1.72a (0.28) 1.47a (0.22)
Freeze Thaw loss % 12.06a (0.67) 10.39a (0.57)
Treatments followed by the same letter in the rows are not significantly different
(p<0.05)
When the data were analysed for differences between consumers evaluation of
venison from carcasses hung by either pelvic suspension or Achilles tendon the pelvic
suspension methods scored significantly higher for tenderness (p=0.004) and juiciness
(p=0.011). (Table 4)
Table 4: Mean sensory evaluation scores for venison from fallow deer hung by either
the Achilles tendon or by pelvic suspension
Hanging Achilles Hung Pelvic Suspension
Colour 8.74a (0.18) 8.09b (0.19)
Aroma 8.39a (0.16) 8.80a (0.15)
Aroma Strength 7.80a (0.17) 7.46a (0.18)
Flavour 9.74a (0.16) 9.91a (0.19)
Flavour Strength 8.52a (0.16) 8.25a (0.19)
Game Flavour 6.71a (0.20) 6.75a (0.21)
Tenderness 8.82a (0.21) 9.64b (0.19)
238 R
Hanging Achilles Hung Pelvic Suspension
Juiciness 7.64a (0.21) 8.39b (0.20)
Overall Liking 9.92a (0.18) 10.38a (0.20)
Treatments followed by the same letter in the rows are not significantly different
(p<0.05)
Red deer carcasses hung by pelvic suspension were preferred by panelists for
tenderness (p<0.001), juiciness (p=0.001) and overall liking (p=0.007). (Table 5)
Table 5: Mean sensory evaluation scores for venison from red deer hung by either the
Achilles tendon or by pelvic suspension
Hanging Achilles Hung Pelvic Suspension
Colour 8.37a (0.27) 7.93a (0.29)
Aroma 8.98a (0.24) 8.90a (0.24)
Aroma Strength 7.52a (0.27) 7.46a (0.28)
Flavour 9.97a (0.25) 10.36a (0.25)
Flavour Strength 8.01a (0.24) 8.22a (0.24)
Game Flavour 6.58a (0.31) 6.57a (0.30)
Tenderness 8.87a (0.33) 10.85b (0.24)
Juiciness 8.90a (0.29) 10.22b (0.29)
Overall Liking 10.37a (0.28) 11.38b (0.24)
Measurements in columns with the same later are not significantly different
The technique to hang carcasses by the pelvic bone (obturator foramen) instead of
the normal position by the Achilles tendon resulted in more tender meat in the M.
Longissimus dorsi for fallow deer bucks, does and red deer stags; determined by shear
force measurements and consumer preference. It is well known that the conditions
during rigor development (e.g. muscle pH decline, temperature/pH relationship and
carcass treatment) are very important in controlling meat tenderisation (Dransfield,
1994). Therefore, carcass suspension techniques have been studied for beef ( Hostetler
et al., 1970: Lundesjo Ahnstrom et al., 2003) where variation in tenderness is
considered to be the main reason for consumer dissatisfaction (Koohmaraie, 1996).
The positive effect of pelvic suspension on tenderness in venison from fallow and red
R 239
deer in this study is important information to consider for the Australian deer industry.
In addition, the important commercial cuts from female deer were generally more
tender than the same cuts from males. The slaughter of female deer therefore provides
a good option for farmers wishing to supply chilled venison year-round, especially at
times of the year when the quality of venison from male deer may be affected by the
breeding season. Given the consistency of this result in this and other studies (Sims
et al 2004) and the importance of meat tenderness in the meat retail sector (MSA,
2001) this technique should be adopted by the deer industry. Inconsistency of product
appears to be a major difficulty in establishing repeat purchasing of venison by
consumers in Australia (Cox et al, 2006). Equalising meat quality characteristics and
sensory evaluations using a technique such as pelvic suspension should bring about
greater consistency of product.
For this study, consumers clearly distinguished their preference for venison
derived from carcasses treated with pelvic suspension post slaughter, with the
important characteristics of tenderness and juiciness improved by this technique. This
finding is consistent with the biochemical data and indicates that pelvic suspension
should be adopted by the deer industry to produce venison that consumers have an
increased preference for and to bring about more consistency in product quality. Use
of this post slaughter carcass treatment is likely to provide significant return on
investment because of the anticipated improvements to product quality and
consistency. The need to adopt such processes in carcasses of all ages, sexes and
condition is unequivocal if enhanced tenderness and juiciness of venison is desirable.
The sensory panels in this study validated the biochemical tests. The technique for
pelvic suspension can be easily installed as routine practice in abattoirs by altering the
mechanics of how a carcass is manipulated on the meat rail in and out of the chiller.
Carcasses can be rehung by the Achilles tendon for ease of transportation once a core
body temperature below 7ยฐC is reached.
References
Cox, R, Watson K & McCrae, T (2006) The Australian Vension Industry. RIRDC,
ACT, Australia.
Dransfield, E.(1994). Optimization of tenderisation, ageing and tenderness. Meat
Science 36:105-121.
Hostetler, R. L., Landmann, W. A., Link, B. A. & Fitzhugh Jr, H. A. (1970). Influence
on carcass position during rigor mortis on tenderness of beef muscles: comparison of
two treatments. Animal Science 31, 47-50.
Koohmaraie, M. (1996). Biochemical factors regulating the toughening and
tenderisation processes of meat. Meat Science 43:193-201.
Lundesjรถ Ahnstrรถm, M., Enfรคlt, L., Johansson, J., Virhammar, K., Hansson, I.,
Johansson, L. & Lundstrรถm, K. (2003). Effect of pelvic suspension on sensory and
instrumental evaluation on four beef muscles in heifers and young bulls. In
Proceedings 49th International Congress of Meat Science and Technology, Sao Paolo,
Brazil, 161-162.
240 R
Meat Standards Australia. (2001). How to do it. Beef CRC, Armidale, NSW,
Australia.
SPSS (2002) SPSS for Windows Version 11.5, SPSS Inc. Chicago, Illinois, USA.
Mulley, R.C, Hutchison, C.L., Flesch, J.S, Wiklund, E, Nicetic, O (2006) The
Relationship of Body Condition Score with Consuemr Perception of Venison Quality.
RIRDC,ACT, Australia.
Sims, K.L., Wiklund, E., Hutchison, C.L., Mulley, R.C. and Littlejohn, R.P. (2004)
Effect of Pelvic Suspension on the tenderness of meat from fallow deer (Dama dama).
Proc. 50th Int. Congress of Meat Science and Technology, Helsinki, Finland.
Wiklund, E., Pickova, J., Sampels, S., & Lundstrรถm, K. (2001) Fatty acid composition
in M. longissimus lumborum, ultimate muscle pH values and carcass parameters in
reindeer (Rangifer tarandus tarandus L) grazed on natural pasture or fed a commercial
feed mixture. Meat Science 58:293-298.
(Oral presentation.)
192
Contents of toxic metals (Cd, Pb, Hg) in tissues of the red deer
(Cervus elaphus) living in the wild.
A. Dobrowolska and K. Gรณrecka
Department of Animal Anatomy, Agricultural University of Szczecin, Poland
Dynamic development of industry and transport, increasing use of chemicals in the
agriculture, and emergence of large urban agglomerations resulted in constantly
growing accumulation of toxic metals such as mercury, cadmium, and lead in various
components of the natural environment. The threat posed by accumulation of those
metals stems from their high toxicity and increasing environmental loads. Toxic
effects of the metals on living organisms have resulted in the necessity of monitoring
metal levels in the environment and in animal tissues. The amount of toxic metals
absorbed by an organism depends on numerous factors, including the extent of
environmental contamination, physical and chemical properties of compounds that
include the metals, and on animal age and physiological condition. Moreover, toxic
metals are a serious problems related to game meat consumption, as game animals
contribute significantly to human nutrition in many countries. The study was aimed
at determining contents of toxic metals in tissues of wild red deer and to determine the
resultant threat, at prolonged exposures, to humans and animals. Analyses were
performed on organs (kidneys and liver) and muscle and nervous tissue collected from
25 red deer shot in north-western Poland in the 2005/2006 hunting season. The organs
(livers, kidneys) as well as samples of the muscle and nervous tissue (the latter from
the telencephalon)were collected directly after the animals were shot by hunters and
R 241
were analysed for the content of cadmium (Cd), mercury (Hg), and lead (Pb). The
cadmium contents found were high, as opposed to those of lead and mercury. The
highest cadmium contents were found in kidneys (3.212.19 mg/kg wet weight). The
mean liver and muscle cadmium contents were 0.310.19 and 0.150.09 mg/kg wet
weight. The lowest mean cadmium content (0.130.05 mg/kg wet weight) was recorded
in the brain tissue, but the content was highly in excess of the admissible level. The
high cadmium content in the red deer may be a result of environmental contamination,
as the animals assayed were shot in an area featuring a large power station and its
waste dumps.
(Poster presentation.)
193
Variations in characteristics of fat, free amino acids and taste of
meat of Japanese deer.
M. Ishida
1
, T. Inoue
1
, T. Mashiko
2
, K. Souma
2
, and S. Ikeda
3
1
Miyagi University, 2-2-1 Hatatate, Taihaku, Sendai 982-0215, Japan
2
Tokyo University of Agriculture, Japan
3
Miyagi Agricultural College, Japan
Variations in characteristics of fat, free amino acids and taste of meat of Japanese deer
reared at three different farms are reported in this study. Comparisons were made
among; a doe fed lucern hay cube, beat pulp and barley (deer 1) and another doe fed
these with additional egoma seeds (perilla seeds) (deer 2), both reared for 50 days at
farm A; four each of stag and doe fed tofu residue and beat pulp (deer 3) and three
does fed grass hay with dried sea weed (deer 4) reared for 14 months at farm B; two
stags fed grass hay and lucern hay cube (deer 5) for 5 months at farm C. Total lipid
contents in loin were 3.81%, 6.86%, 4.91%, 3.19% and 2.89% for deer 1,2,3,4 and 5,
respectively. In fatty acid composition, levels of palmitic acid were higher (P<0.01)
for the deer of farm B than those of other farms while the deer of farm A had a much
higher (P<0.01) mean alfa-linolenic acid content (2.31%) due to adding egoma seeds
to the diet compared with other deer (0.56% and 0.79% for farm B and C,
respectively). The deer of farm C had a higher mean level of conjugated linoleic acid
than that of farm A (0.18mg/g vs 0.02mg/g). For free amino acids, the deer of farm A
and B tended to have elevated levels of amino acids related to bitterness than those of
farm C. The deer of farm C had higher (P<0.05) levels of glutamic acid and lower
(P<0.01) levels of leucine and they also obtained higher scores for tenderness and
strength of taste by a sensory panel while other deer were regarded to be juicier, fatter
and better overall. It is noted that the higher the levels of palmitic or palmitoleic acids
the stronger the flavour or the better overall taste. The meat having higher levels of
stearic and linolenic acids tended to have reduced flavour.
(Poster presentation.)
242 R
Deer zooarcheology and
history
244 R
194
Stable isotopes evidence of seasonality effects on diet and locomotor
adaptations of Pleistocene deer from southern Spain.
J. A. Estรฉvez
1, 2, 3
, A. Grandal-dโAnglade
4
, T. Landete-Castillejos
1, 2, 3
, A. J. Garcรญa
1, 2,
3
, and
.
L. Gallego
1, 2
1
Departamento de Ciencia y Tecnologรญa Agroforestal, ETSIA, Universidad de
Castilla-La Mancha. 02071 Abacete, Spain.
2
Secciรณn de Recursos Cinegรฉticos, IDR, Universidad de Castilla-La Mancha. 02071
Albacete, Spain.
3
Instituto de Investigaciรณn en Recursos Cinegรฉticos, IREC (CSIC, UCLM, JCCM).
Campus Universitario s/n, 02071 Albacete, Spain.
4
Instituto Universitario de Xeoloxรญa, Universidade da Coruรฑa.
Campus de Elviรฑa 15071 A Coruรฑa, Spain
Studies of feeding strategies using stable isotope distributions in skeletal tissues
are well known (Cerling and Sharp, 1996). In fact, stable isotope records (
13
C/
12
C and
18
O/
16
O) are powerful, established tools for reconstructing paleodiets and
paleoenvironmental conditions (Bocherens et al., 1994; Zazzo et al., 2002). By
observing seasonal variation in isotopic ratios, we should be able to asses whether
specific forage (browse or graze) is always preferred, or whether a diet fluctuates
between browsing and grazing conditions. By using ฮด
13
C ratios in animal tissues, and
since the isotopic ratio of the ingested plant will be there recorded, we can infer the
photosynthetic pathway of plants, and therefore, vegetation type. Several factors might
explain variations in ฮด
18
O values of carbonate hydroxylapatite, but water ingested and
leaf water, in consequence the diet, should play a prominent role in causing different
ranges in ฮด
18
O ratios. These values of animal tissues can be used to infer the ฮด
18
O of
ingested water, which is in turn related to that of climate and precipitation, although
leaf water can be isotopically modified relative to precipitation due to evaporation at
the leaf surface (Fricke et al., 1998). In this study, water sources may have included
the nearby lagoons, fruits and leaves of terrestrial plants. Mostly browsers are drought
tolerant species, and, on the other hand, grazers use to be obligate drinkers (Cormie
and Schwarcz, 1996). Seasonal temperature changes can also be assessed by analyzing
oxygen isotope variation. In mid to high latitude continental areas, there is a strong
positive correlation between ฮด
18
O ratios and temperature. Warmer temperatures would
result in a heavier ฮด
18
O ratio, while cooler temperatures would result in lighter ฮด
18
O
values.
From a morphological point of view, feeding strategies can be established based
on craniodental characteristics. Mammalian herbivores with short-crowned
(brachyodont) teeth are usually assigned as predominantly browsers, which mainly
consume leaves and fruits of trees and shrubs in forested environments or woodlands
(e.g. Eucladoceros giulii and Stephanorhinus etruscus). On the other hand,
mammalian herbivores (e.g. Equus altidens and Bovini aff. Leptobos) with high-
crowned (hypsodont) teeth are classified as predominantly grazers, which are defined
as consumers that feed primarily, or exclusively, on abrasive grasses or other low
R 245
herbs typically associated with more open grassland biomes (MacFadden, 1997;
Palmqvist et al., 2003).
In the study here presented, we have further re-examined existing carbonate
hydroxylapatite
13
C/
12
C and
18
O/
16
O data variability in four different large bodied
species of mammal herbivores (Stephanorhinus etruscus, Eucladoceros giulii, Equus
altidens and Bovini aff. Leptobos) from the fossil rich early Pleistocene (ca. 1.3Ma)
assemblage of Venta Micena in the Guadix-Baza Basin, southern Spain (Palmqvist et
al., 2003). In this lithological section, vertebrate remains have been rapidly deposited
in a swampy/lacustrine sequence, under savanna type climate conditions (Palmqvist
et al., 2003). Our aim was to compare the stable isotopic ratios and dental morphology
in order to characterize diet and feeding behavior. In principle, these species exhibit
two different dietary adaptations: browsers and grazers feeders (Tab. 1).
Table 1. Published feeding strategies, mean ฮด
13
C and ฮด
18
O values, standard errors of
the mean, and number of sampled individuals. Data from Palmqvist et al., 2003.
ฮด
13
Cฮด
18
O
Specimen Codea Mean S.E.Mb. n Mean S.E.Mb. n
Equus
altidens
G -7.7 0.2 12 -3.1 0.2 12
Bovini aff.
Leptobos
G -7.5 0.3 8 -3.3 0.2 8
Eucladoceros
giulii
B -6.9 0.2 8 -3.6 0.2 8
Stephanorhin
us etruscus
B -7.5 0.2 5 -4.2 0.2 5
a
G: grazers; B: browsers.
b
S.E.M.: standard error of the mean.
We have found patterns that can be interpreted in terms of diet due to vegetational
and seasonal climatic changes. Isotope ฮด
13
C values in carbonate hydroxylapatite have
a mean of -7.1ยฑ0.1โฐ for browsers (n=20) and โ7.6ยฑ0.2โฐ for grazers (n=13),
suggesting a predominant C
3
feeding diet, mainly composed by trees, shrubs and
grasses from relatively cool/humid climates (Tab. 1 and Fig. 1). If the skeletal tissue
has ฮด
13
C values that are less than -10โฐ then animals have consumed C
3
plants; if ฮด
13
C
values are more than -2โฐ, then they have feed C
4
plants (Lee-Thorp and van der
Merwe, 1987). More specifically, vertebrate hervibores with ฮด
13
C values lower than
-8โฐ has been described as pure C
3
feeders, requiring no C
4
contribution to the diet
(Cerling et al., 1997). However, in a C
3
-dominated region, it is rather difficult to
distinguish between browsers and grazers only based on ฮด
13
C values (Zazzo et al.,
2002).
246 R
Figure 1. ฮด
13
C and ฮด
18
O values in carbonate hydroxylapatite of herbivores from Venta
Micena (southern Spain). Data from Palmqvist et al., 2003.
The ฮด
18
O values from browsers range from -4.8 to -2.8โฐ, and values from grazers
are at -4.3 to -2.2โฐ (Fig. 1). Nevertheless, if lighter (winter) ฮด
18
O values characterized
by a first quartile of -4.0โฐ are similar among the browsers and grazers, the heavier
(summer) ฮด
18
O ratios (third quartile -2.9โฐ) are only present in three out of the four
species (E. giulii, E. altidens and Bovini aff. Leptobos). Two major factors affect the
isotopic composition of leaves: the isotopic composition of the soil water and the
degree of evaporative enrichment of this water after it reaches the leaves. The effect
of leaf water is greatest during summer months since evaporative enrichment is
affected by humidity and temperature. Grasses, which are abundant during this season
in savanna environments, are often highly enriched in
18
O as a result of the
evapotranspiration. Therefore, and under these conditions, the isotopic composition
of a diet rich on grasses should be distinguished by heavier ฮด
18
O values, as we have
observed in measurements from grazers. Although C
3
plants constitute about 90% of
all plants, today 80 to 85% of the savanna-grass species in tropical and temperate
climates are C
4
plants, and only less than 10% are C
3
. These are predominantly found
near closed forests (Schwartz, 1991). Based on this, we suggest that the environment
at Venta Micena was mainly characterized by an open savanna woodland, rather than
by an unforested savanna (Palmqvist et al., 2003).
Eucladoceros giulii is an extinct cervid with a body mass ca. 300kg and
magnificent antlers conformed by many irregularly branched tines. Despite its dental
morphology, isotope data comparisons between browsers and grazers suggest a
feeding strategy for E. giulii different to the already described. The similarity with
grazers could be due to the fact that this species may tend to be a grazer rather than
a browser during the summer, with high herbage mass availability. Therefore, this
R 247
large deer should probably be a mixed feeder, like most of modern deer, especially
during the summer when body and ornamentation size forced them to live in more
open habitats, instead of inhabiting forests. Feeding adaptations and locomotion
strategy of the species depend on the landscape features and body weight. We suggest
that E. giulii was probably adapted to relatively open environments. It has been
proposed that deer ingest grasses in areas with relatively high summer temperatures,
although switching to a diet based on grasses, with potentially lower water content,
could lead to water stress, since deer obtain most of the water necessities from leaves
(Cormie and Schwarcz, 1996). Consequently, although dental characters have been
frequently considered to determine feeding preferences, tooth variability can be
difficult and controversial to interpret as a dietary proxy.
References
Bocherens, H., Fizet, M., Mariotti, A. (1994) Diet, physiology and ecology of fossil
mammals as inferred from stable carbon and nitrogen isotope biogeochemistry -
implications for Pleistocene bears. Palaeogeography, Palaeoclimatology,
Palaeoecology 107(3-4):213-225.
Cerling, T.E., Sharp, Z.D. (1996) Stable carbon and oxygen isotope analysis of fossil
tooth enamel using laser ablation. Palaeogeography, Palaeoclimatology,
Palaeoecology 126(1-2):173-186.
Cerling, T.E., Harris, J.M., MacFadden, B.J., Leakey, M.G., Quade, J., Eisenmann,
V., Ehleringer, J.R. (1997) Global vegetation change through the Miocene/Pliocene
boundary. Nature 389:153-158.
Cormie, A.B., Schwarcz, H.P. (1996) Effects of climate on deer bone delta N-15 and
delta C-13: Lack of precipitation effects on delta N-15 for animals consuming low
amounts of C-4 plants. Geochimica et Cosmochimica Acta 60(21):4161-4166.
Fricke, H.C., Clyde, W.C., O'Neil, J.R. (1998) Intra-tooth variations in delta O-18
(PO
4
) of mammalian tooth enamel as a record of seasonal variations in continental
climate variables. Geochimica et Cosmochimica Acta 62(11):1839-1850.
Lee-Thorp, J., van der Merwe, N.J. (1987) Carbon isotope analysis of fossil bone
apatite. South African Journal of Science 83:712-715.
MacFadden, B.J. (1997) Origin and evolution of the grazing guild in New World
terrestrial mammals. Trends in Ecology & Evolution 12(5):182-187.
Palmqvist, P., Grokke, D.R., Arribas, A., Farina, R.A. (2003) Paleoecological
reconstruction of a lower Pleistocene large mammal community using biogeochemical
(delta C-13, delta N-15, delta O-18, Sr : Zn) and ecomorphological approaches.
Paleobiology 29(2):205-229.
Schwartz, D. (1991) Measurement of delta C-13 as a pedological and ecological
marker of the savanna-forest relationship in equatorial ecosystems. Cahiers-
248 R
ORSTOM, Serie Pรฉdologie 26(4):327โ341.
Zazzo, A., Mariotti, A., Lecuyer, C., Heintz, E. (2002) Intra-tooth isotope variations
in late Miocene bovid enamel from Afghanistan: paleobiological, taphonomic, and
climatic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 186(1-2):
145-161.
195
Biometry and palaeoecology of the Red deer (Cervus elaphus Linnรฉ,
1758) during middle and upper Pleistocene in Western Europe. The
example of the Lazaret cave (Alpes-Maritimes; France)
M. Liouville
1
, P. Valensi
2
, and E. Psathi
2
1
Institut de Palรฉontologie Humaine, Equipe d'archรฉozoologie, 1, rue Renรฉ Panhard,
Paris, France
2
Laboratoire Dรฉpartemental du Lazaret 33 bis, boulevard Franck Pilate, Nice, France
In order to characterize the morphological variations of the red deer during middle and
upper Pleistocene, we intend to establish a link between biometrical data and a
palaeocological approach. On the biometrical point of view, we are using, for the first
time with the red deer, the methodological approach of J. Weinstock, the V.S.I.
(Variability Size Index). The environmental reconstitution of the middle where the red
deer evolved is made possible by the use of several complementary palaeocological
methods. Even if we are aware of their limits, we try to understand our results through
a naturalist but careful approach. The second part of our doctoral research is to
describe, for each studied site, the palaeoethnological aspect of the relationship
between humans and deer. The zooarcheaological approach also allows us to estimate
the impact of the human factor within the bone accumulation, using this aspect of their
behaviour to illustrate the potential changes towards hunting. The site of the Lazaret
cave, with an exceptional stratigraphy, allows us to examine the case of a
pre-Neanderthal group. Moreover, the interdisciplinary analysis undertaken since the
beginning of the excavation, give us the opportunity to approach more closely the
everyday life of this disappeared specie.
(Oral presentation.)
R 249
196
Fallow deer of Rhodes: an ongoing, comprehensive study about
ecology, genetics and conservation.
D. Mertzanidou
1
and A. Legakis
2
1
Department of Biology, University of Athens, Rhodes, Greece
2
Aethrea: AgroEnvironmental Research and Action Team, Greece
Unlike other parts of the world where fallow deer are widespread due to human
activity, and treated as a park animal or pest on some occasions, fallow deer has had
a distinct historical course on the island of Rhodes, in Greece. The fallow deer
population of Rhodes is considered to be the only free-ranging population of ancient
origin that survived on a Mediterranean island, and the only wild population in
Greece. Besides the great research interest that it has presented (as an insular
population living in a vulnerable Mediterranean ecosystem), its severe population
decrease in late 1980s - early 90s and its recognition as a protected species by Greek
law, as well as its cultural and symbolic value for the island of Rhodes pushed the
need for thorough research to be conducted to identify and apply appropriate
conservation practices. Three years ago the first systematic recording of past and
present fallow deer distribution and habitat was begun, and progressively was enriched
by data about fallow deer feeding ecology and genetic structure. Recently, a pilot
telemetry study has been being conducted, and conservation measures, such as the
creation of a compensation system for damages caused by fallow deer on summer
crops, are in process. The current presentation is actually an activity report, which
aims at giving a comprehensive picture of fallow deer of Rhodes through the
objectives, the methods, some results and the future goals of the ongoing study.
(Poster presentation.)
250 R
Contributions received and
accepted after the deadline
252 R
197
Conservation of huemul (Hippocamelus bisulcus) deer in Chilean
Patagonia: a new research initiative.
P. Corti
Dรฉpartement de Biologie, Universitรฉ de Sherbrooke, Sherbrooke, Quรฉbec, Canada
The huemul or southern Andean deer is a flagship species of the mountainous and
forested habitats of Chile, principally in the southern Patagonian region. It is listed by
the IUCN as critically endangered and it has declined to under 2000 animals. Possible
proposed causes of this decline are habitat loss, poaching, diseases and competition
from livestock, and predation. However, we just begin to understand the ecology of
huemul and much more need to be done. The reality is that nothing is known about
metapopulation structure and connectivity, recruitment rates and survival of juveniles,
population dynamics and genetic diversity, and the limiting factors that negatively
affects the growth of remaining populations. These uncovered important issues for
huemul conservation motivated this new research in southern Chile, located at ''Lago
Cochrane National Reserve'', Aysรฉn District.
The goals of the study are to determine possible limiting factors for huemul population
persistence and growth. The initial actions include the increment of the amount of
marked animals by a previous project in order to be more accurate with our
measurements of population sizes, survival rates, ranging behaviour, patterns of
habitat use, population genetics, and social organization. Huemul present an unusual
social organization for a mid-sized sexually dimorphic ungulate, forming small mixed
groups with high site fidelity of both sexes that would make them more susceptible to
fail reproduction. However, we know little about the relationship between the mating
system of huemul and the variation in demographic parameters, such as population
density. Mating strategies are studied through behavioural observations of known
individually marked animals and corroborated through modern genetic tools to study
genetic relationships and hence mating successes in the study huemul populati ons.
We are also tagging newborn deer to determine survival of young animals and
recruitment into the adult cohort, this action allows us to know the whole life of an
individual animal since is born. The ultimate goal is to provide relevant quantitative
data to improve management strategies for the Chilean government through CONAF
and the IUCN/SSC Deer Specialist Group for the conservation of Hippocamelus
bisulcus in Chile.
(Poster presentation.)
R 253
198
Translocation and semi-captive breeding of huemul (Hippocamelus
bisulcus) with purpose of reintroduction in Chile.
P. Corti
Dรฉpartement de Biologie, Universitรฉ de Sherbrooke, Sherbrooke, Quรฉbec, Canada
The huemul has experienced a strong reduction in its distribution in Chile and
Argentina and its number is no more than 2000 individuals. Due to this grave situation
the Huilo-Huilo Foundation, a Chilean NGO, planed the creation of a huemul breeding
centre. The main goal of this initiative was the reintroduction in the western side of
the Andes range in southern Chile (39ยบS), which is part of a private protected area, the
Huilo-Huilo Biological Reserve. We planed to extract a unit of six huemul (1 adult
male, 1 juvenile male, 2 adult females, and 2 juvenile females) to form a founder
population and to prevent a large impact on donating populations. The extraction area
was the Aysรฉn District (Patagonia), which has the healthiest and largest huemul
populations. We selected places where huemul survival is critical because of the
presence of domestic livestock and dogs, roads, outside protected areas were poaching
is stronger. The first 2 deer (1 male and 1 female) were captured in late April. The
adult couple were chemically immobilized, accommodated in boxes, and awaked with
a reversal agent to be transported via helicopter and airplane. Each deer was measured
and health checked, radio-collared, and sampled. The couple was released in a 64 ha
enclosure of suitable habitat after 4 and half hours of flying. This initiative is a
landmark for conservation in South America, but generated controversy among local
political authorities. The major of local village set a demand against the Foundation
and SAG (Chilean Agricultural Service), and the project was stopped. In June 2005
the judge failed in favour of the Foundation. Today governmental authorities are
conditioning the initiative through several requirement and restrictions.
In December 2005 we recorded the first offspring born in semi-captivity for this
project. In February 2006 we added another adult female taken by SAG from a peasant
that illegally kept it. Actually, the 4 animals are in excellent conditions and we are
expecting 2 fawns to be added in late November.
(Poster presentation.)
254 R
199
So similar and yet so different: The surprising polyphyletic origin
the genus Mazama (Mammalia: Cervidae).
J. M. B. Duarte
1
, S. Gonzรกlez
2
, and J. E. Maldonado
3
1
Depto Zootecnia, FCAV/UNESP, CEP 14884900 JaboticabalSP, Brazil.
2
Depto. GenรฉticaIIBCEFacultad de Ciencias/UdelaR Av. Italia 3318 Montevideo
11600 Uruguay
3
Genetics Program, National Zoological Park/National Museum of Natural History,
Smithsonian Institution, 3001 Connecticut Ave. NW, Washington D.C. 20008, USA
The systematic relationships of the extant Neotropical deer and their evolutionary
history in South America have been poorly studied. We undertook a molecular genetic
analysis using cytochrome b gene sequences of all representative genera of neotropical
deer in order to resolve the controversial systematic relationships among this group
of New World deer species. Here we combine morphological and cytogenetic data and
find that morphologically conservative species show high levels of molecular and
cytogenetic divergence. Our phylogenetic analysis revealed that brocket deer of the
genus Mazama have undergone an amazing morphological convergent evolution with
a highly divergent phylogenetic history. The evolutionary history of this group began
with various ancestral forms in North America that gave rise to red brockets, gray
brockets, marsh deer, pampas deer, pudu, huemul, and white tailed deer that entered
South America via the Panama land bridge (2.4 MYA) and evolved separately but the
ancestral form of the red brocket group continued an explosive speciation process
generating various independent forms (M. nana, M. bororo, M. americana, and M.
temama). The high levels of genetic divergence between gray and red brockets
suggests a polyphyletic origin. Furthermore, we found such high levels of genetic
divergence between Mazama nemorivaga and M. gouazoubira that we recommend
that they be placed in different genera (Coassus and Cariacus). The example we
present here describes one of the most amazing cases of morphological convergent
evolution in mammals where brocket deer with very similar morphologies show high
levels of molecular and chromosome differentiation and evolutionary diversification.
(Oral presentation.)
R 255
200
Factors affecting the composition of autumn diet of red deer (Cervus
elaphus) in Alpine environment
M. Heroldovรก
1
, M. Homolka
1
, J. Kamler
1
, C. Ghezzi
2
, W. Redaelli
3
, E. Andreoli
2
, and
S. Mattiello
2
1
Institute of Vertebrate Biology, Brno, Czech Republic
2
Istituto di Zootecnica, Faculty of Veterinary Medicine, University of Milan, Via
Celoria 10, 20133 Milan, Italy
3
Hunting Management Committee, Sondrio, Italy
The effect of sex, lactation stage, date and place of culling on the autumn diet
composition of red deer in Central Italian Alps (Val Fontana, Province of Sondrio)
was evaluated by analysis of vegetal fragments in rumen content of 47 individuals.
Percentage volume (% v), percentage frequency (% f) and the relative importance
index [I
i
= (%f
i
+ %v
i
)/2] were estimated for each food item. Trophic diversity (Hยด)
and equitability index (Jยด) were calculated. Grasses were the most important
components of deer diet in terms of volume (41% v), followed by shoots and leaves
from broadleaved species (28% v). Herbs (10% v, 28% f, I = 19%) and Rubus sp. (8%
v; 4% f, I = 6%) were additional important components of deer diet, and some items
of ferns, seeds and dwarf shrubs species were also found in the rumens. Only 4 %v of
the diet was represented by concentrated food (i.e. seeds and fruits). In spite of the
prevalence of grasses, diet diversity (Hยด = 2.63) and equitability (Jยด = 0.56) were high,
probably because of the wide altitudinal range used by red deer in this alpine valley
(from 700 to above 2200 metres a.s.l.). Neither sex nor lactation stage significantly
affected diet composition. Significant differences were found in response to the date
of culling: in late autumn (October-November) deer made a higher use of grasses and
a lower use of broadleaved and Rubus species than in early autumn (September),
probably in response to the progressive fall of leaves from the trees, which induced
the deer to be more grazers than browsers (early autumn: 35% v of grasses, 31% v of
broadleaved species, 11% v of Rubus sp.; late autumn: 57% v of grasses, 18% v of
broadleaved species, 1% v of Rubus sp.). Place of culling seemed to affect diet
composition: deer harvested at higher elevations (above 1940 m a.s.l.; 6 individuals)
showed a higher presence of grasses (74% v in average, max. 90%). These results
confirm the high level of adaptation of red deer to different environmental conditions
and to the related changes in food availability.
256 R
Author index
รcs
, Z.
......................78
Adams, K. A. ................119
Affandy, L. .................. 187
Agungpriyono, S. ............. 189
ร
hman, B............... 26, 30, 32
Alarcos, S............ 138, 197, 213
Aldridge, D. .................106
Allen, S. .................... 161
Althnaian, T. .................161
Ando, M. ................ 154, 219
Andreoli, E................ 81, 255
Apollonio, M. ......... 17, 130, 131
Archunan, G. .................111
Arnold, W. .................. 173
Attinault, K. .................125
Audenaerde, P. M. F. ..........164
Ayanegui-Alcรฉrreca, M. A. .... 70, 71
Bรกdr, V....................... 82
Balfanz, F.................... 173
Baranฤekovรก, M. ............ 41, 50
Barbosa, A. M. ................56
Barbosa, J.....................56
Bar-David, S. ......... 23, 105, 128
Barna, R. .................... 157
Barrio, J. .................... 172
Bartoลก, L. . . . 132, 133, 136, 137, 139,
163, 164, 169
Bartoลกovรก-Vรญchovรก, J. . 132, 133, 136,
137, 139
Beck, R....................... 79
Beiglbรถck, C. ................173
Bello, J. ......................47
Bello-Gutiรฉrrez, J...............55
Berndt, A. ....................56
Berruecos, J. M. ...............92
Bertoletti, I. ...................81
Bertolotto, E..................131
Beszterda, P. ..................24
Bianchi, A. ................... 81
Birรณ, Z. ................... 22, 48
Bleier, N............... 22, 40, 212
Bobek, B. ................ 24, 144
Bocci, A. .................... 125
Bona, F...................... 154
Bongi, P. .................... 131
Borkowski, J. .................41
Bowman, J. L................. 119
Braga, G. G. .............. 96, 211
Broekhuijse, M. L. J. W.......... 80
Bubenik, G. A. .... 16, 163, 164, 169
Buฤko, J. .................... 222
Buenrostr, A................... 45
Bunod, A.-H. .................. 38
Caboni, A. ................... 166
Cagnacci, F. ............. 140, 155
Callovi, I. ................... 155
Campbell, J. R. .................4
Campbell, T. A................ 192
Carnevali, L. ..................53
Carr, S. M.....................85
258 R
Carranza, J......... 56, 89, 138, 197
Carriรณn, D. .................. 167
Casagrande, S. ................ 88
Castillo, L. ............... 89, 197
Castillo-Alcala, F. .....70-72, 75, 76
Cavallaro, A. ................. 90
Ceacero, F. .................. 167
Chandola, S. ................. 101
Chapman, N. G. .............. 169
Chapple, D. G. ............ 35, 179
Chen, M..................... 107
Ciuti, S. ................ 130, 131
Clark, H..................... 180
Collins-Emerson, J. M. ....... 70, 71
Conner, M. C. ................ 119
Cooper, S. M. ................. 44
Corona, N.................... 47
Corti, P. ................ 252, 253
Cosse, M. .................... 87
Cรดtรฉ, S. D. .......... 194, 198, 199
Cottrill, B. .................. 179
Coulombe, M.-L. ............. 198
Csรกnyi, S. ................... 152
D'Angelo, G. J................ 193
D'Angelo, J. G................ 193
Danell, ร.................. 26, 30
Daniels, M. J. ................ 151
Davies, M. H. ............. 35, 179
Dayan, T. .................... 23
de Lisle, G. W. ............. 67, 68
De Marinis, A. M. ......... 59, 117
Delgadillo, A. C. .............. 92
Demarais, S. ................. 115
Demiaszkiewicz, A. ............ 47
DeYoung, R. V. .......... 115, 192
Di Luzio, P. ................. 225
Dmuchowski, B............. 46, 47
Dobrowolska, A. ......... 168, 240
Dolev, A. ................ 23, 105
Dominguez-Rebolledo, A. E. .... 120
Drรกbkovรก, J. ............. 133, 139
Draisma, M. ................. 116
Dryden, G. M. ............ 37, 229
Duarte, J. M. B. ........... 56, 254
Duลกek, A. ....... 132, 133, 136, 139
Egri, B....................... 78
Equihua, M. .................. 47
Erdenbaatar, J. ................ 73
Ernst, M...................... 97
Esteso, M. C. ................ 120
Estรฉvez, J. A. ............ 167, 244
Fernรกndez-Garcรญa, J. L. ......... 89
Fรฉrnandez-Santos, M. R. ....... 120
Ferreira, A. J. ........... 22, 42, 53
Filli, F. ..................... 156
Finฤo, S..................... 222
Fischer, A. .................. 129
Flesch, J. S. ................. 234
Fletcher, T. J. ................ 228
Florijanฤiฤ, T.................. 79
Flueck, W. T. ............ 174, 181
Focardi, S. .......... 117, 145, 225
Fodor, ร. ................... 212
Folkow, L. .................. 224
Forchhammer, M. C. ............ 2
Ford, W. M. ................. 192
Formaggioni, P. ............... 51
Frฤ
ckowiak, W. .............. 144
Franceschi, P. ................. 51
Franzetti, B. ................. 145
Frey, R...................... 114
Friฤovรก, B. .................. 137
Fritsch, G.................... 114
Fukuta, K. .................... 91
Furlanello, C. ................ 140
Gallagher, G. R. .............. 193
Gallego, L. .............. 167, 244
Gallina, S. ................. 45, 47
Galvan, B. ................... 87
Gandolfi, G. .................. 88
Garcia, A. J. ................. 167
Garcรญa, A. J. ................. 244
Garde, J. J. .................. 120
Gaspar-Lรณpez, E. ............. 167
Gawor, M. .................. 144
Gazzolo, C. .................. 216
Gebler, A.................... 114
Gee, K. L.................... 115
Geist, V. ...................... 2
Gerkema, M. P. .............. 224
Gerwing, V. ............... 73, 86
Ghezzi, C. ................... 255
Giczi, E. ..................... 78
Gill, R. ..................... 106
R 259
Giลผejewski, Z.................114
Glossop, J. C. ................. 66
Goda, R. .................... 154
Gonzales, R. A................ 115
Gonzรกlez, S. ........... 87, 96, 254
Gรณrecka, K............... 168, 240
Goyal, S. P. ..................210
Gozzi, C. ..................... 59
Grace, N. D. ............... 75, 76
Grandal-dโAnglade, A. .........244
Griffin, J. F. T. ............. 67, 68
Grignolio, S. ................. 131
Gunsch, H. .................. 156
Haanes, H.....................84
Haigh, J. C. ............. 29, 73, 86
Hamann, J.-L. ................ 220
Handarini, R.............. 187, 189
Harrison, W. M. .............. 116
Hata, H.......................52
Hayakawa, D. ........ 121, 123, 124
Heikkilรค, R...................200
Heim, M......................43
Heinzl, E. .................... 81
Hendrichs, H. ................129
Heroldovรก, M. ............. 41, 255
Heuer, C................ 66, 70, 71
Hewitt, D. G. ................. 118
Homolka, M. ........... 41, 50, 255
Honeycutt, R. L. ..............115
Hoskin, S. O...... 38, 69, 72, 80, 180
Huber, S. .................... 173
Huot, J. ......... 194, 195, 198, 199
Hutchison, C. L. .............. 234
Igota, H. ................ 121, 123
Ikeda, S. .................... 241
Illmann, G. .................. 136
Ilyas, O...................... 208
Imperio, S.................... 117
Inoue, T. .................... 241
Ishida, M. ................... 241
Janicki, Z. ....................79
Jarnemo, A. ................... 49
Jiang, G. .................... 108
Jiang, Z...................... 219
Jiggins, C. G. I. ................84
Jirsa, A. ..................... 138
Johnsingh, A. J. T. .............39
Kaji, K. ..................... 153
Kaleem, A. .................. 100
Kamler, J. ................ 41, 255
Katona, K........... 22, 40, 48, 212
Keay, M. G.............. 29, 73, 86
Khan, J. A. .................. 100
Khursheed, A. ................ 104
Kierdorf, H............ 74, 160, 162
Kierdorf, U............... 160, 162
Kimura, J. ....................91
Kissell Jr., R. E. ..............146
Kitamura, N. ......... 121, 123, 124
Kjellander, P. .................54
Klein, F. .................... 220
Kol, N. V. .................... 95
Kolecki, M. ..................144
Kondo, S. ........ 52, 121, 123, 124
Konjeviฤ, D. ..................79
Koprowski, H.................114
Kotrba, R. . . . 133, 136, 139, 163, 169
Kovรกcs, A..................73, 77
Kovรกcs, S. .................73, 77
Kovรกcs, V....................212
Krzywiลski, A. ................46
Kลกรกda, V.....................136
Kumรณr, K. ................... 218
Kuลพmovรก, E. .................169
La Morgia, V. ................ 154
Landete-Castillejos, T. ..... 167, 244
Lanna, D. P. D. ................ 56
Laseter, B. R. ................192
Lazebny, O. E. ................95
Le Corre, M. .................178
Legakis, A. ..................249
Legendre, X. ................. 109
Li, C. ........................18
Lingle, S................. 134, 135
Liouville, M. .................248
Liu, Z. S. .................... 110
Locatelli, Y. .................109
Lohmann, C. H................ 165
Lรณpez, R......................92
Losos, S. .................... 164
Luccarini, S. ................. 130
Luna, A. .....................92
Lรผtjens, I. ...................163
Ma, J. ......................108
260 R
Mackintosh, C. G. .... 67, 68, 70, 71
MacMillan, D. C. .............. 25
Malacarne, M. ............. 51, 88
Maldonado, J. E. ........ 87, 96, 254
Malins, M. I................... 36
Mamok, T.................... 24
Mandal, A. .................. 210
Marasco, V. .................. 59
Mariani, P. ................ 51, 88
Marshall, H. D................. 85
Martรญnez, J. G. ................ 89
Martinez-Pastor, F............. 120
Mashiko, T. ................. 241
Mason, P. .................... 69
Massรฉ, A. ................... 199
Masseti, M.................... 90
Masyud, B. .................. 187
Mateos, C. .......... 138, 197, 213
Matias, D.................... 120
Mรกtrai, K.................. 22, 40
Matsuura, Y....... 52, 121, 123, 124
Mattiello, S. .............. 81, 255
Mattioli, S. .................. 166
Maxwell, C. .................. 65
McGonnell, I. M. ............. 161
McLeod, J. ................... 84
McShea, W. E. ................ 19
Meek, M. G................... 44
Merino, M. L.................. 87
Mermillod, P. ................ 109
Merta, D. ............... 144, 218
Mertzanidou, D. .............. 249
Midwinter, A. C. ........... 70, 71
Mikoล, J...................... 24
Miller, B. F. ................. 192
Miller, K. V. ......... 136, 192, 193
Mishra, S.................... 210
Monaco, A................... 145
Monaco, E. L................. 118
Montaldo, H. H. ............... 92
Montanaro, P................. 225
Moore, G. I. ................. 116
Moore, I. A. ................. 116
Morales, J. M. ............... 152
Moreira, M. Z. ................ 56
Motro, U. ................... 209
Mount, J. G. ................. 161
Muller, L. I. ................. 119
Mulley, R. C. ................ 234
Murgia, C. .................. 166
Muzylak, M.................. 161
Mwendwa, J. M................ 80
Nagy, S. ................. 58, 122
Nalley, W. M. M. ......... 187, 189
Nansalmaa, M. ................ 73
Napp, J. ................ 160, 162
Nasiadka, P. .................. 41
Neteler, M. .................. 140
Niittee, E. .................... 57
Niลผnikowski, R. ............... 47
Nishitani, K................... 52
Nugent, G. ................... 66
Nuรฑez, A. M. ................ 217
Nyeste, M. .................. 212
Nygrรฉn, K. .................. 114
Okushima, S. ................ 161
Olajos, T. ................... 212
Oliveira, A. M. ................ 42
Olofsson, A. .................. 26
Ondris, M. ................ 38, 69
Orosz, S................... 22, 40
Osborn, D. A. ................ 193
Otsuka, S.................... 123
Owens, M. K. ................. 44
Pados, Z...................... 58
Palme, R. ................... 173
Panamรก, J. .............. 136, 137
Panasiewicz, G. .............. 114
Pecchioli, E. .................. 90
Pedrotti, L. ...... 140, 145, 155, 156
Pellerin, M................... 178
Pellis, S. M. ............. 134, 135
Pemberton, J. ................. 84
Pรฉntek, I. ................... 122
Pรฉnzes, Z..................... 78
Perelberg, A............... 23, 105
Perez-Barberia, F. J............. 84
Perez-Espona, S. ............... 84
Pรฉrez-Gonzรกlez, J. ......... 56, 197
Persson, I. L. ................ 196
Piasentier, E................... 51
Pintur, K. .................... 79
Pisani, G. M. .............. 51, 88
Pluhรกฤek, J. ......... 133, 136, 139
R 261
Pokhrel, S. ................... 102
Pomroy, W. E............ 69, 72, 80
Potvin, F..................... 195
Prasad, S. N. ..................39
Preisler, J. .................... 82
Price, J. S. ...................161
Prokeลกovรก, J. ............... 41, 50
Psathi, E. .................... 248
Purwantara, B............. 187, 189
Puskรกs, E. ................... 122
Putzu, N. ....................154
Qureshi, Q. ..................104
Raimondi, V. ..................87
Rajagopal, T. ................. 111
Ramalho, R. M................. 53
Randveer, T. ..................57
Rawat, G. S. .................. 39
Redaelli, W. .................255
Reimoser, F. ................. 210
Rembacz, W. ..................24
Rendall, D. .................. 135
Richards, A. ..................74
Richards, E. D. ................85
Richter, H.....................74
Riga, F. ......................53
Rรธed, K. H. ................84, 86
Rolf, H. J. ....... 160, 162, 163, 165
Roll, U. ..................... 105
Ronchi, F. ................... 117
Rosef, O. ..................... 84
Sabbioni, A. ..................88
Said, S. ..................... 178
Saikkonen, T. ................ 196
Saint-Andrieux, C. ............ 220
Saltz, D........... 23, 105, 128, 209
Sรกnchez-Prieto, C. B. . . 138, 197, 213
Sรกnchez-Rojas, G...............45
Sasaki, M. ........... 121, 123, 124
Sathyakumar, S. ........... 39, 104
Sato, H. ..................... 154
Saucedo, C. ..................106
Sรฆther, B.-E...................43
Scanziani, E. ..................81
Schams, D. .................. 164
Schliephake, H............ 160, 162
Semiadi, G. ..................187
Severin, K. ................... 79
Seymour, N. ............. 160, 162
Sharma, C. P. ................210
Shibata, E................ 154, 219
Shorthose, W. R............... 229
ล iler, J. .....................164
Silva, C....................... 22
Simard, A.................... 194
Singh, R. R. .................. 210
Sinha, B. C...................101
Sinha, S. P. .................. 101
Skuterud, L....................32
Slavica, A.....................79
ล mรญdovรก, E. .............. 136, 137
Smith-Flueck, J. M...... 85, 174, 181
Snochowski, M. ...............46
Soffiantini, C. S. ............ 51, 88
Solberg, E..................... 43
Song, Y.-L. .................. 105
Sookhareea, R. ...............229
Souma, K. ................... 241
Standio, A. ................... 24
Starz, J. ......................47
Steplewski, Z. ................ 114
Steyaert, S. .................. 222
ล tindl, P. ..................... 82
Stokkan, K. A. ................ 224
Storms, D. B. F. .............. 220
Subandriyo .................. 189
Subekti, K. .................. 187
Sugรกr, L. ........... 73, 77, 78, 157
Summer, A.................... 51
Suominen, O. ................196
ล ustr, P.................. 136-138
Sutama, I. K. ................. 187
Suter, W. .................... 223
Suttie, J. M. ...................18
Suzuki, M......... 52, 121, 123, 124
ล vecovรก, L. .......... 132, 133, 139
Swainson, N. M. .............. 180
Szabรณ, J. ..................... 58
Szafranska, B. ................ 114
Szรฉkely, J. ...................212
Szemethy, L. ........ 22, 40, 48, 212
Tagliabรฒ, A. .................155
Tagliavini, J. .................. 88
Takahashi, H. ................153
Takayanagi, A. ............... 186
262 R
Tappe, P. A. ................. 146
Tatsuzawa, S. ................ 226
Tedford, C.................... 64
Telford, M................... 125
Terhes, A.................... 212
Thapa, T. B. ............. 102, 103
Thompson, J. ................. 68
Toelihere, M. R. .......... 187, 189
Tomรกnek, M. ................ 169
Torres, J..................... 197
Toso, S. .................. 53, 59
Tremblay, J.-P. ............ 43, 195
Tsubota, T. .................. 121
Tume, R..................... 229
Tyler, N. J. C................. 224
Ueda, K. ..................... 52
Urbano, F. .................. 140
Valencia, J............... 138, 197
Valensi, P. .................. 248
Vallet, J.-C. ................. 109
Van Laere, G................. 178
van Oort, B. E. H.............. 224
Vaลkovรก-Formanovรก, D. ....... 136
Vรกsquez, G. C. ................ 92
Vernesi, C. ................... 90
Wang, X. M.................. 110
Wappel, A. L.................. 44
Warren, R. J. ................ 193
Wasilewski, R. ................ 24
Weissengruber, G. E. .......... 114
Weladji, R. B................. 194
Whelan, K. J. ................. 37
Widmer, O................... 178
Wiese, K. G.......... 160, 162, 163
Wilhelm, J.-L. ............... 220
Wilson, P. R. . . 38, 62, 63, 66, 69-72,
75, 76, 80
Wilson, W. F. ................ 134
Wiลniowska, L................ 144
Woodbury, M. R. ............... 4
Wu, J. ...................... 111
Yanagawa, Y...... 52, 121, 123, 124
Yayota, C. ....... 52, 121, 123, 124
Yusuf, T. L. ................. 187
Zakharov, I. A. ................ 95
Zeng, Z. .................... 105
Zhang, D.-X. ................ 105
Zhang, E. ................... 107
Zhang, M.................... 108
Zhang, Q. ................... 105
Zhang, Y. ................... 111
Zidon, R. ................... 209
Zink, R. .................... 210
Zomborszky, Z. ........... 58, 122
Zweifel-Schielly, B. ........... 223
Index
behaviour
cooperative behaviour .......136
browsing
long term chronic browsing . . . 194
mating strategies
alternative mating strategies....17
absolute population density ........24
abundance .... 25, 50, 53, 55, 87, 100,
102-104, 110, 145, 150, 154-
156, 195, 196, 199
overabundance ......25, 191, 197
activity budget .............198, 199
Africa ..................5, 19, 166
aggression ................112, 133
agonistic behaviour .........111, 177
agriculture . . ii, iv, 5, 8, 13, 22, 48, 52,
56, 64, 97, 107, 121, 123, 124,
146, 153, 179, 186, 210, 229,
232, 233, 240, 241
agricultural production ........19
agricultural technology .......5, 9
Alceini........................85
Alces alces .......7, 43, 54, 200, 204
allosucking ...............132, 133
America . iii, 2-5, 7, 15, 16, 64, 85, 172,
182, 253, 254
North America . 2-4, 7, 15, 64, 254
South America ...... 16, 85, 172,
182, 253, 254
ammonia .................179, 180
analysis software
Computer-Assisted Semen
Analyzers ............122
ISAMUD .................140
animals . . . 5-15, 22-24, 27, 29, 30, 33,
35, 43, 46, 47, 51, 54-56, 58,
59, 63, 64, 66-69, 73, 76, 80,
81, 86, 91, 92, 95-97, 100,
107, 114, 121, 122, 128, 129,
133, 138, 140, 144, 150, 152,
155, 157, 166, 169, 179-181,
184, 185, 187, 192, 193, 200,
203, 218, 223, 226, 229, 235,
240, 241, 245, 247, 252, 253
wild animals ................10
annual cycle ..................224
anti-predatory behaviour .........136
antler
antler composition ..........167
antler cycle ................189
antler growth . . 18, 117, 164, 169,
187
antler size .........175, 176, 197
antler structure .............168
Haversian canals............168
interstitial lamellae ..........168
pedicle periosteum .......18, 161
264 R
Volkmannโs canals....... 51, 53, 94,
118, 179, 189, 229-231, 235-
237, 241
antlers . . . 18, 49, 85, 86, 101, 116, 117,
139, 152, 158, 160-169, 187,
211, 246
apoptosis ..................... 165
artificial barriers............... 213
artificial feeding ........ 4, 10, 12, 31
Asia . . . iii, 5, 13, 16, 19, 91, 210, 211,
232
China . . 5, 105, 107, 108, 110, 111
India . . . iv, 39, 100-102, 104, 111,
208, 210, 211
Israel ..... 23, 105, 106, 128, 209
Mongolia ........ 29, 73, 86, 111
assisted reproduction
artificial insemination . . . 122, 188
Australia....5, 37, 116, 229, 234, 239,
240
authochtonous population ......... 88
Axis
Axis axis ................... 10
Axis kuhlii ................ 187
axis deer
bawean deer............... 187
balanced sex ratio.............. 119
barasingha ................... 100
bark ..................... 50, 219
bark stripping .............. 50, 219
barking ......... 5, 39, 102, 182, 211
barking deer ......... 5, 39, 102, 211
behavior . 5, 7, 129, 174, 175, 177, 178,
182, 184, 185, 193, 194, 198-
200, 245
behaviour .......22, 38, 40, 106, 111,
112, 116, 117, 125, 127, 129,
131-133, 135, 136, 139, 140,
175-178, 222, 225, 229, 248,
252
altruistic behaviour ......... 135
foraging behavior........... 198
reproductive behaviour ...... 116
spatial behavior ............ 178
biogeography .............. 85, 172
birth . 43, 46, 55, 92-94, 114, 131-133,
139, 167, 212
birth weight .............. 132, 133
Blastocerus
Blastocerus dichotomus ...... 172
body condition . 26-28, 31, 53, 72, 118,
194, 219, 234, 240
body mass . . . 30, 31, 43, 138, 157, 173,
176, 194, 246
body size ....27, 28, 39, 45, 133, 166,
167, 180, 197
body temperature . . 148, 150, 235, 239
body weight . 27, 30, 31, 46, 47, 55, 75,
91, 114, 167, 188, 247
bone marrow fat ................ 53
Bos taurus .................... 52
Bot fly
Cephenemyia auribarbis ...... 77
Pharyngomyia picta .......... 77
bottleneck ..................... 86
brain .................. 73, 75, 241
breeding lifespan .............. 116
breeding season . 44, 58, 109, 123, 125,
174, 176, 239
brocket deer
grey brocket deer............ 56
browsing . . . 44, 50, 192, 194-196, 198,
200-204, 210, 212, 213, 218,
244
browsing impact .... 50, 213, 218
Brucella
Brucella abortus ............ 81
Brucella ovis ............... 62
Caesium ................... 32-34
calf . . . 27, 28, 30, 31, 36, 73, 94, 132-
134, 139, 140, 182
calving ............... 43, 117, 131
Canis latrans ............. 134, 135
Canis lupus ................... 204
Capreolus . . . 22, 54, 70, 74, 78, 85, 88,
136, 165, 178, 204, 220
Capreolus capreolus 22, 54, 70, 74, 88,
136, 165, 178, 204, 220
Capreolus capreolus italicus ... 54
carbohydrate .......... 179, 187, 188
carbon................... 179, 247
carcass . . . 26-28, 30-32, 67, 232, 234,
238-240
R 265
cattle . . . 4-8, 11, 13, 15, 52, 67, 92, 94,
97, 179-181, 183, 212, 232,
233
cementum annuli ................59
central nervous system ..........183
central vessels .................165
Cervidae ...... 42, 53, 57, 58, 78, 85,
172, 232, 254
Axis ................10, 11, 187
Odocoileinae ...............85
Odocoileus . . . 6, 8, 10, 12, 13, 15,
44, 45, 70, 85, 86, 115, 119,
134-136, 146, 172, 194, 195,
198, 217, 233
Odocoleinae ...............116
cervids . . 2, 4-10, 12, 14, 16, 56, 65, 78,
85, 182, 184, 194, 196, 219
elk....... 2-19, 21-25, 29, 35-59,
61-82, 84-94, 96, 97, 99-111,
114-125, 128-140, 144-158,
160-169, 171-182, 184-187,
191-200, 203, 204, 208-214,
217-226, 228-230, 232-234,
237-241, 243, 244, 247-249,
252-255
huemul ....85, 106, 107, 181-185,
252-254
moose . . . 7, 29, 43, 54, 57, 58, 66,
196, 200, 201, 203, 204
taruca .....................85
Cervinae
Alces alces ...............4, 12
Dama . . 10, 23, 47, 51, 54, 59, 70,
78, 90, 105, 117, 128, 130,
131, 136, 137, 160, 162, 163,
165, 209, 225, 233, 234, 240
Pudu ...........16, 86, 172, 254
Cervus . 6, 7, 10, 11, 19, 22, 24, 37, 39,
40, 46, 48, 49, 52-54, 57, 58,
67, 68, 70, 71, 73, 74, 77, 78,
81, 84, 89, 91, 92, 94, 97, 100,
101, 103-105, 108, 109, 114,
116, 120-124, 132, 133, 136,
138, 139, 144, 145, 151-154,
156, 162-166, 168, 173, 174,
178, 187, 189, 197, 219, 220,
222, 223, 226, 229, 232-234,
240, 248, 255
Cervus duvauceli .......100, 101
Cervus elaphus . . . 6, 7, 10, 11, 22,
24, 37, 40, 46, 48, 49, 53, 54,
57, 58, 67, 68, 70, 71, 73, 74,
77, 78, 81, 84, 89, 92, 94, 97,
104, 108, 114, 120, 122, 132,
133, 136, 138, 139, 144, 145,
151, 152, 156, 162-164, 166,
168, 173, 174, 197, 220, 222,
223, 233, 234, 240, 248, 255
Cervus eldi .............19, 105
Cervus timorensis . . 187, 189, 229,
233
Cervus duvauceli
Cervus duvauceli duvauceli . . 100,
101
Cervus elaphus
Cervus elaphus atlanticus .....84
Cervus elaphus canadensis ....97
Cervus elaphus corsicanus ....166
Cervus elaphus hanglu .......104
Cervus elaphus hippelaphus ... 46,
58, 122
Cervus elaphus hispanicus 89, 94,
120, 138
Cervus elaphus manitobensis ....7
Cervus elaphus nannodes ......11
Cervus elaphus scoticus .......92
Cervus elaphus xanthopygus . . 108
Cervus eldi
Cervus eldi hainanus ........105
Cervus nippon . . 52, 91, 109, 121, 123,
124, 153, 154, 219, 226, 232
Cervus nippon yesoensis ......52
Cervus unicolor .............39, 116
Cervus unicolor unicolor .....116
Chernobyl accident ...........32, 34
chromatin ....................120
chromosome ..................254
climate . . 3, 4, 16, 22, 48, 53, 107, 157,
173, 174, 223, 225, 244, 245,
247
commercial shooting .............26
community . . 29, 54, 63, 86, 101, 186,
195, 247
invertebrate communities .....196
266 R
vegetation communities ...... 198
compensation ......... 132, 133, 249
competition . . . 17, 54, 57, 58, 130, 252
Intersexual competition ....... 17
conception ................ 80, 117
condition . . 2, 22, 23, 26-28, 30, 31, 44,
53, 57, 58, 67, 72, 118, 157,
182, 187, 188, 194, 203, 219,
234, 239, 240
conservation ....2, 3, 6, 13, 19, 25, 50,
54, 65, 87, 90, 96, 100-103,
105-109, 111, 155, 184, 197,
207, 208, 210, 212, 213, 216,
249, 252, 253
wildlife trade ........... 14, 210
contraception
Gossypol ................. 114
cooperation ................... 210
corpora lutea.................. 121
countryside ................. 36, 37
culling ..... 11, 25, 26, 36, 59, 68, 255
culture . . . 12, 29, 62, 67, 68, 70, 71, 74,
109, 160, 162, 163, 232
cytochrome b gene .... 85, 87, 96, 254
dam......................... 136
Dama
Dama mesopotamica 23, 105, 128,
209
Dama dama . 10, 47, 51, 54, 59, 70, 78,
90, 117, 130, 131, 136, 137,
160, 162, 163, 209, 225, 233,
234, 240
day length.................... 224
deer.... 4, 6-8, 11-13, 16, 44, 45, 115,
116, 119, 134-136, 146-151,
182, 192-195, 198, 199, 233
axis deer................ 10, 11
barasingha ................ 120
barking deer...... 5, 39, 102, 211
brocket deer...... 55, 56, 86, 254
caribou.... 26, 28, 43, 66, 95, 224
Chinese water deer ..... 107, 108
fallow deer . . 10, 11, 17, 23, 47, 51,
54, 59, 78, 90, 105, 106, 117,
128, 130, 131, 136, 137, 160,
162, 163, 165, 166, 177, 178,
209, 225, 233, 234, 237, 238,
240, 249
Hangul ................... 104
hog deer .......... 101-103, 211
lesser mouse deer............ 91
maral ..................... 46
marsh deer ............. 85, 254
mule deer . . 8-10, 14, 15, 134, 135,
182, 204
non-seasonal deer .......... 221
pampas deer . . 85, 87, 96, 211, 212,
254
roe deer ............ 30, 31, 233
rusa ......187-189, 229-231, 233
sambar deer ............... 116
seasonal deer .............. 221
sika deer....52, 91, 109, 121, 123,
124, 153, 154, 219, 226, 232
wapiti.....10, 19, 48, 56, 74, 101,
103, 107, 108, 110, 120, 122,
148, 149, 175, 182, 216, 235,
236, 244, 246, 247
deer farming . . 4, 5, 35, 36, 58, 63, 180,
233
deer- forest systems ............ 195
deer-vehicle collisions ...... 193, 194
density estimation
aerial detection rate ......... 148
censusing..... 143, 144, 150, 155
distance sampling . . 145, 147-150,
155, 156
fecal-pellet group count ...... 154
relative density index........ 144
thermal infrared imaging . 146-149
dentine ............ 74, 75, 138, 139
fluorotic coronal dentine ...... 74
dentine depletion .............. 138
diagnosis ........... 68, 72, 167, 184
diel rhythmicity ............... 224
diet .....22, 31, 32, 35, 37, 38, 40, 41,
45, 46, 50, 54, 55, 110, 111,
118, 167, 180, 181, 186, 194,
216, 220, 223, 227, 230, 241,
244-247, 255
diet composition . . 22, 37, 40, 110,
223, 255
digestion..................... 179
R 267
disease . 4-15, 36, 62-68, 70, 71, 73, 81,
182-184
Bovine tuberculosis . . . 4, 6, 7, 10,
13, 15, 36, 68, 69
chronic wasting disease . 4, 6, 8, 9,
13-15, 62-66
Elaphostrongylus cervi ........73
fascioloidosis ...............78
fluorosis ...................74
Johneโs disease . . . 4, 6, 10-12, 67,
68
Leptospira............70, 71, 81
leptospiral shedding ..........62
parasitic bronchopneumonia ....81
sarcosporidiosis .............81
serovars .............70, 71, 81
spongiform encephalopathy . . 114,
120, 122, 188
tuberculosis . 4, 6-8, 10, 11, 13-15,
36, 62, 63, 67-69
yersiniosis..................63
zoonotic disease ..............5
dispersal ...........84, 85, 192, 209
sex biased dispersal ..........84
distribution . 16, 22, 27, 35, 43, 44, 53,
55-58, 76, 84, 85, 88, 89, 100,
102, 104, 106-109, 128, 155,
156, 165, 172, 174, 177, 197,
200, 204, 208, 209, 213, 214,
216, 217, 225, 235, 249, 253
spatial distribution . . 56, 108, 213,
214
spatio-temporal distribution ....44
DNA . 6, 74, 78, 86-89, 91, 95-97, 105,
120, 192, 193, 211
cDNA .....................18
rDNA .....................78
doe..........101, 118, 119, 182, 241
domestic animals.... 35, 92, 114, 180,
184
goat...................12, 185
sheep . 8, 16, 22, 26, 28, 32, 44, 45,
47, 48, 58, 77, 81, 88, 109,
114, 118, 120-123, 125, 131,
138, 173, 174, 176, 177, 179,
197, 216, 218, 223, 233, 239,
240, 246
domestication ...............29, 95
dominance....111, 112, 116, 125, 177
dominance hierarchy ....111, 112
dominance hierarchy formation
................111, 112
dose.......................68, 72
dummy ......................136
ecology . . iii, 2, 4, 5, 10-13, 23, 24, 28,
40, 41, 55, 100, 104, 106, 128,
144, 166, 169, 173, 178, 185,
192, 196, 209, 210, 213, 215,
216, 218, 220, 247, 249, 252
autecology ..................2
Ecosystem . . . 3-5, 7, 8, 10-13, 34,
50, 177, 196, 204, 216, 249
Habitat . . . 6, 8, 12, 17, 19, 22, 39,
42-44, 47-50, 53-55, 65, 87,
88, 96, 100-103, 106-111,
119, 130, 131, 138, 153, 155-
157, 172-175, 177-179, 181,
183, 184, 187, 196, 197, 199,
200, 203, 204, 208, 209, 211-
213, 217, 219, 220, 223, 225,
226, 249, 252, 253
ecotone ..............174, 177, 213
embryo ......................109
England......................228
environment . . 2, 3, 5, 7, 9-11, 22, 35,
36, 40, 41, 57, 65, 66, 92, 105,
118, 131, 140, 152, 179, 199,
209, 224, 225, 240, 246, 255
lowland . . . 19, 22, 35, 36, 103, 172
mountain . . 4, 7, 13, 15, 22, 30, 84,
111, 145, 223
woodland . . . 36, 37, 106, 228, 246
environmental contamination . . 10, 12,
66, 240, 241
epidemiology . 4, 62, 65, 66, 68, 70, 71,
81
eradication.............7, 10, 65, 66
estrus........................124
Europe....iii, 3, 16, 41, 74, 78, 84, 88,
90, 177, 197, 210, 248
Austria ............78, 173, 210
Croatia ....................79
Czech Republic ......... 41, 50,
74, 78, 82, 89, 97, 114, 122,
268 R
129, 132, 133, 136-139, 160,
162-165, 169, 185, 186, 211,
220, 255
Denmark ................ 2, 74
Estonia................. 57, 58
Fennoscandia ............... 54
Finland....31, 115, 196, 200, 201,
204, 205, 240
France ....94, 109, 122, 125, 178,
179, 220, 248
Greece ................ 90, 249
Hungary .......22, 40, 48, 58, 73,
77, 78, 122, 152, 157, 212
Italy . 17, 51, 53, 54, 59, 81, 88, 90,
117, 125, 130, 131, 140, 145,
154-156, 166, 225, 255
Norway .... 31-34, 41, 43, 84, 86,
89, 224
Poland ....24, 25, 41, 46, 47, 114,
144, 168, 218, 240
Portugal ............. 22, 42, 53
Russia .................... 95
Spain . . . 56, 77, 89, 120, 138, 166,
167, 197, 213, 244-246
Sweden .... 26, 30, 32-34, 49, 54,
196
evolution....3, 22, 23, 83, 85, 91, 115,
128, 135, 178, 195, 209, 224,
228, 233, 247, 254
evolutionary radiation ........... 85
extinction . . 8, 104, 109, 156, 186, 208,
212
faeces . 33, 41, 47, 76, 77, 79, 173, 209
fallow deer
Persian fallow deer . 23, 105, 106,
128, 209
farming . . 4, 5, 9, 10, 12, 35, 36, 58, 63,
66, 109, 180, 213, 233
farmed deer . 6, 36, 38, 62, 64, 66-
68, 70-72, 76, 80, 179, 180
fatness score................... 27
fatty acids................ 229-233
fawn ....100, 101, 104, 115, 116, 118,
121, 134, 135, 182
fawning................... 48, 119
feeding
feeding ecology .... 40, 215, 216,
249
feeding strategies .......... 244, 245
female . . . 17, 27, 28, 30, 31, 43, 46, 52,
56, 57, 59, 82, 85, 94, 100,
104, 111, 114-117, 119, 121,
128, 130, 131, 133, 134, 137,
157, 174-176, 178, 179, 182,
235, 239, 253
female choice ............. 117, 137
female defence ................ 134
femaleโs reproductive state ....... 135
fetus ........................ 183
fir saplings ................... 218
food . . . 1-5, 7-9, 13, 26, 31, 32, 35-37,
40, 41, 45, 51, 55, 57, 58, 63,
64, 91, 100, 103, 110, 111,
114, 131, 153, 167, 173, 177-
179, 182, 186, 199, 201, 203,
212, 219, 220, 222-224, 227-
235, 237-240, 255
food availability ....173, 177, 201,
219, 220, 223, 255
food quality ......... 31, 51, 219
food search ................... 178
foraging ......... 131, 174, 198, 199
forest . . . 7, 8, 22, 24, 25, 30, 36, 39-41,
43, 45, 48-50, 53-55, 57, 97,
101-103, 106, 107, 111, 124,
136, 138, 144-146, 153, 172,
174, 175, 178, 181, 186, 192,
195, 196, 198, 200, 201, 203,
204, 210, 212, 217-220, 222,
223, 225, 226, 247
cloud forest ............... 217
floodplain forest ............. 50
tropical dry forest ........... 45
forest floor ................... 196
free amino acids ............... 241
game....13, 29, 32, 40, 58, 65, 66, 79,
97, 150, 152, 155, 157, 186,
200, 204, 210, 212, 235, 237,
238, 240
gene flow ............... 84, 85, 87
genetic differentiation . . . 87, 105, 193
genetic diversity ........ 90, 107, 252
genetic relatedness ............. 193
genetic variation......... 84-87, 105
R 269
genetics . . . 17, 83, 84, 87, 92, 95, 130,
131, 249, 252
conservation genetics .........87
genotype ...................69, 78
gestation . . 16, 48, 109, 116, 117, 121,
183, 219
giant river fluke
Fascioloides magna .......78, 79
glands ....................91, 123
caudal gland ...............123
metatarsal gland ............123
pre-orbital gland............132
salivary glands ..............91
global environmental changes.......2
North Atlantic Oscillation . . . 4, 7,
104, 137, 138, 145, 156, 210,
213, 216
glucocorticoids
glucocorticoid metabolites ....173
GPS . . 44, 58, 119, 138, 140, 146, 147,
152, 179, 199, 223
grandmother ..................134
grassland . 100-103, 107, 110, 175, 226,
245
grazing . . 8, 30, 31, 33, 35, 37, 38, 76,
81, 101, 136, 196, 212, 213,
222, 244, 247
growth . . 18, 23, 28, 31, 38, 62, 67, 75,
80, 92-94, 104, 107, 114, 117,
128, 157, 161-164, 169, 173,
181, 182, 184, 187, 195, 218,
224, 229, 233, 252
fetal differentiation..........164
IGF-1 .................86, 119
insulin-like growth factor I....169
proliferation .......162, 163, 169
habitat
habitat heterogeneity ......47, 48
habitat quality . . 157, 173, 200, 204
habitat segregation ..........226
habitat selection .... 42, 110, 111,
131, 138, 199, 203, 223, 225
habitat use . . . 39, 43, 44, 100, 103,
138, 172, 199, 200, 204, 208,
211, 213, 217, 220, 225, 252
habitat choice ...............49, 54
habitat fragmentation . 55, 87, 108, 109
habitat selection
scale-dependent habitat selection
....................223
Hangul
Kashmir Stag ..........104, 211
haplotype...................87, 89
harem . . . 115, 125, 137, 174-178, 213
harem defense......174, 177, 178
harvest . . . 25, 49, 53-55, 119, 144, 153
herbivore . . 40, 41, 138, 181, 184, 212,
220
herbivores . . 41, 50, 54, 179, 184, 197,
198, 213, 220, 244-246
herbivory.................192, 213
herd . . . 6, 7, 12, 14, 26, 30, 51, 64, 67,
71, 73, 86, 92, 112, 115, 119,
133, 134, 177, 192, 193
heterozygosity ..................95
hind . . . 36, 57, 94, 109, 116, 133, 134,
139, 140, 157, 233
Hippocamelus
Hippocamelus antisensis 172, 216,
217
Hippocamelus bisulcus . . 106, 181,
252, 253
histology .....................168
histopathology...............68, 74
history . . iii, 2-4, 11, 43, 53, 60, 82, 87,
116, 119, 182, 194, 228, 243,
254
evolutionary history .....182, 254
phylogenetic history .........254
home range . . 22, 23, 43, 47-49, 54, 86,
117, 119, 131, 138, 140, 157,
178, 222, 225
home range size..........48, 54, 138
hominids .....................228
hormone ..........16, 123, 124, 164
cortisol ...............164, 173
follicle stimulating hormone ....16
glucocorticosteroids .........173
luteinizing hormone ..........16
melatonin ..............16, 164
prolactin ...............16, 164
steroids production ..........121
testosterone..... 16, 162-164, 169
hormones
270 R
dihydrotestosterone ..... 162, 163
human..... 3-7, 10-12, 17, 32, 50, 55,
62, 65, 103, 107, 110, 130,
148, 208, 212, 213, 226, 228,
240, 248, 249
human activities ..... 4-7, 10, 12, 103
hunting . . 4, 7, 8, 12, 24, 25, 29, 40,
53, 55, 58, 79, 81, 88, 106,
107, 120, 140, 144, 146, 156,
157, 181, 184, 186, 192, 197,
208, 212, 220, 228, 240, 248
hunters . . . 8, 25, 29, 36, 186, 210, 240
hunting
dogs ....... 5, 107, 182, 184, 253
overhunting ............... 184
hurricane Lothar............... 220
hybrids ....................... 46
Hydropotes ................ 85, 107
Hydropotes inermis ......... 107
Hydropotes inermis inermis ... 107
hypothesis . . . 12, 17, 18, 58, 130, 132-
134, 136, 137, 161, 165, 181,
185
compensation hypothesis .... 132,
133
forage selection hypothesis . . . 17,
130
indirect competition hypothesis
................. 17, 130
predation risk hypothesis . . 17, 130
Indian blackbuck .............. 111
Antelope cervicapra ......... 111
Indonesia ................ 187, 189
industry .....5, 9, 10, 37, 62, 179, 180,
239, 240
introduction . . . 4, 7, 28, 53, 57, 65, 92,
174, 181, 200, 209
invertebrate communities ........ 196
invertebrates.................. 196
ISAMUD
Information System for Analysis
and Management of
Ungulates Data . . . 145, 194
IUCN ....... 107, 109, 208, 210, 252
IUCN's Red data list............ 104
jaw length.................. 27, 46
karyotype ..................... 55
kidney . 53, 62, 70, 71, 75, 81, 229-231
kidney fat index ................ 53
kin selection .................. 135
laryngeal air sac ............... 114
Lazaret cave .................. 248
legislation............... 2, 36, 111
lek.......................... 117
lekking ........... 17, 117, 177, 178
leptospirosis .......... 62, 70, 71, 81
life history . . . 3, 43, 60, 116, 119, 194
lifespan.............. 116, 138, 139
liver .... 75, 76, 78, 79, 235, 240, 241
liver fluke
American liver fluke ......... 79
Fasciola hepatica ........... 79
giant liver fluke ............. 78
livestock . . . 4-13, 36, 64, 81, 180-182,
184, 230, 252, 253
liveweight.................. 69, 80
liveweight gain................. 80
lungworm .................. 69, 78
Dictyocaulus ............... 78
lynx ......................... 29
male ....16, 24, 27, 46, 49, 59, 82, 94,
100, 109, 112, 114-117, 119,
120, 133, 137, 138, 152, 165,
175-177, 208, 233, 235, 239,
253
mallin ................... 175-177
management . . . 4, 6, 8, 10, 15, 19, 21-
26, 28, 29, 31, 32, 34, 36, 37,
40-42, 47-50, 52-54, 56, 57,
62, 63, 65-67, 70, 87, 88, 90,
97, 99, 103, 105, 115, 118,
138, 140, 145, 146, 150, 152,
155-157, 167, 178, 181, 185,
186, 192, 193, 197, 200, 204,
210, 212, 213, 216-219, 225,
229, 234, 252, 255
clear-cuts ............. 195, 198
management strategies . 53, 62, 70,
87, 192, 252
monitoring . . 26, 32, 33, 43, 49, 65,
72, 79, 81, 100, 102, 104, 156,
181, 185, 194, 200, 204, 205,
210, 240
R 271
wildlife management..... 4, 6, 10, 28,
41, 42, 146, 150, 210
marsh deer
Blastocerus .............85, 172
mate.............111, 117, 119, 137
mating . . 16, 17, 45, 56, 111, 112, 115-
117, 121, 125, 174, 177, 178,
214, 252
mating season.......45, 121, 177
mating system . . . 56, 116, 214, 252
mating strategies . . . 17, 117, 174, 177,
178, 252
mating success ................125
Mazama .....55, 56, 86, 172, 217, 254
Mazama americana .........172
Mazama bricenii ............217
Mazama gouazoubira .....56, 172
Mazama gouazoubira nemorivaga
.........................172
Mazama nemorivaga ........254
Mazama rufina .............172
Mazama temama ............55
meat.....4, 5, 29-35, 67, 92, 180, 211,
228, 229, 232-241
Mediterranean climate.....22, 53, 225
metabolism . . . 76, 173, 179, 183, 219,
232
methane..................179, 180
microsatellite markers .....84, 97, 193
migration.............7, 16, 30, 222
seasonal migration ..........222
milk....11, 30, 31, 51, 68, 74, 94, 133
mineralization ........16, 74, 75, 164
mitogenic effect............162, 163
molecular phylogeny .............85
morphology ....82, 165, 187, 245, 246
morphometry ...................45
mortality . 10, 107, 150, 153, 157, 183,
195, 201, 218, 226
neonatal mortality ...........183
Moschidae
Moschus ...........39, 111, 208
Moschus berezovskii .........208
Moschus chrysogaster ....39, 208
Moschus fuscus .............208
Moschus moschiferus ....111, 208
mother ................43, 115, 135
mtDNA ............. 88-90, 95, 107
Munitacus
Munitacus muntjac ...........39
Muntiacus reevesi ....91, 129, 169
Muntiacus reevesi micrurus ...129
musk deer .........39, 111, 208, 211
black musk deer ............208
dwarf musk deer............208
Himalayan musk deer.....39, 208
Siberian musk deer......111, 208
Mycobacterium ....4, 6, 10, 14, 15, 67
Mycobacterium avium . . 4, 10, 14,
67
Mycobacterium bovis .....4, 6, 15
Mycobacterium avium subsp.
paratuberculosis .....10, 14
natality ......................226
needles .......................41
neonatal survival...............185
neural crest cells ...............161
New Zealand . . 1-5, 18, 38, 62, 63, 66-
72, 75, 76, 80, 180
nitrogen....45, 51, 179, 194, 229, 247
nitrous oxide ..................179
North America
Canada . iii, iv, 2, 4, 5, 7, 9, 13, 14,
16, 29, 65, 66, 73, 85, 95, 134,
135, 163, 164, 169, 194, 195,
198, 199, 252, 253
Mexico ......9, 45, 47, 48, 55, 92
United States . . 5, 10, 44, 64, 150,
193
North atlantic oscillation ...........3
nursing ..........133, 134, 139, 182
nutrition .... 46, 58, 68, 118, 122, 167,
180, 219, 228, 232, 240
nutritional deficiency ........184
Odocoileinae
Capreolini .................85
Odocoileini .................85
Hippocamelus
........ 85, 106, 172, 181,
216, 217, 252, 253
Odocoileus . . . 6, 8, 12, 13, 15, 44, 70,
115, 119, 134-136, 146, 172,
194, 195, 198, 217, 233
Odocoileus hemionus . . . 8, 15, 134
272 R
Odocoileus viginianus mexicanus
.......................... 45
Odocoileus virginianus clavium
.......................... 12
Odocoileus virginianus peruvianus
.................... 172
oestradiol .................... 169
Oestridae ..................... 77
offspring....17, 31, 68, 107, 109, 116,
131, 133, 135, 253
oocyte....................... 109
ovaries .................. 109, 121
overabundance ...... iii, 25, 191, 197
overgrazing.................... 44
ovulation ................. 125, 194
Ozotoceros ............. 85, 96, 211
Ozotoceros bezoarticus . . . 96, 211
palaeocological methods ........ 248
palaeoethnological aspect ........ 248
pampas deer
Ozotoceros .......... 85, 96, 211
parasites ................ 69, 72, 78
Brucella ............. 62, 74, 81
hemoparasite ............... 74
Nasopharyngeal bot fly ....... 77
parasitism .................. 72, 80
Sub-clinical parasitism ....... 80
parasitology ......... 46, 47, 79, 185
parental care .................. 135
allosucking............ 132, 133
pasture . . 17, 26, 32, 33, 35-38, 46, 51,
57, 69, 76, 80, 130, 136, 180,
229, 231, 240
pathogen ........... 4, 7, 10, 12, 167
pathogen pollution ......... 4, 10, 12
pathogenesis ................... 68
PCR ..... 68, 88, 89, 95, 96, 161, 211
pellet groups . . . 58, 102, 103, 154, 156,
200, 201, 203, 204
Peru ................. 16, 172, 216
phenotype.................... 166
phenotypic variation.............. 3
photic modulation .............. 224
photoperiod ............... 16, 173
photoperiodic control ........... 224
Phthiraptera ................... 82
Linognathidae .............. 82
Phylogeography ................ 89
physiology . 46, 92, 114, 164, 173, 233,
247
plants . 52, 57, 110, 111, 118, 153, 180,
181, 195, 197, 209, 220, 244-
247
Pleistocene . . . 2, 16, 87, 244, 245, 247,
248
Pliocene ............... 16, 87, 247
pollution ............. 4, 10, 12, 179
population ....3, 6-8, 10-12, 14, 22-26,
28, 36, 43, 46, 52-55, 57, 58,
70, 71, 73, 79, 81, 84-88, 90,
95, 96, 100-108, 124, 125,
128, 144, 145, 149-158, 161-
163, 167, 172-174, 176-178,
181, 182, 184-187, 195-198,
200, 203, 204, 208, 209, 212,
213, 218, 220, 226, 228, 249,
252, 253
population density . . 7, 10, 12, 24,
55, 144, 154, 156, 157, 173,
176, 197, 198, 200, 203, 204,
209, 218, 252
population dynamics . . 3, 53, 153,
156, 220, 226, 252
population expansion ...... 23, 85
population density
density estimation ...... 146, 203
populations abundance ...... 145
sustainable population density
.................... 197
population genetic structure . . . 84, 105
population genetics .......... 84, 252
population growth . . 23, 104, 107, 157,
181
population size . . . 24, 57, 96, 100, 105,
128, 145, 152, 153, 182, 200,
203
predation.....2, 17, 29, 107, 118, 130,
131, 134, 135, 182-185, 204,
252
predator
bear .................. 6, 7, 29
coyote ..................... 7
fox ........................ 6
lynx ..................... 124
R 273
wolf ..............29, 186, 204
predators . . . 2, 182, 183, 198, 199, 226
pregnancy .....68, 121, 124, 131, 183
prey .............134, 182, 196, 228
production ....... 5, 7, 19, 30-32, 35,
36, 47, 51, 52, 54, 63, 71, 72,
92, 94, 109, 112, 118, 121,
132, 133, 136, 137, 139, 163,
164, 169, 179, 180, 189, 234
progeny ...................35, 133
progesterone ..............121, 124
protein.... 5, 9, 14, 40, 46, 51, 52, 55,
161, 179, 180, 189, 219, 223
public . . 5, 8, 26, 36, 37, 65, 107, 186,
196
pudu ..............16, 86, 172, 254
Pudu mephistophiles .....16, 172
Southern pudu ..............16
radiation .............23, 32, 35, 85
Radioactivity..........30, 31, 34, 35
radiotracking ...................47
rainfall....................48, 118
range . 11, 22, 23, 28, 39, 43, 44, 47-49,
53, 54, 66, 72, 76, 77, 80, 86,
87, 101, 116, 117, 119, 128,
131, 138, 140, 145, 148, 151,
156, 157, 164, 167, 172, 174-
176, 178, 185, 187, 208, 209,
217, 222, 223, 225, 246, 253,
255
Rangifer . 26, 28, 30-32, 43, 85, 95, 97,
114, 224, 240
Rangifer tarandus . . 26, 28, 30-32,
43, 95, 97, 114, 224, 240
Rangifer tarandus tarandus 30-32,
224, 240
ranging strategy
suboptimal ranging strategy ....48
rank .... 116, 125, 139, 140, 173, 176,
225
receptors .................123, 124
androgen receptor .......123, 165
estrogen receptor .......123, 124
progesterone receptor ........124
reciprocal altruism..............135
recolonization .........181, 182, 193
red brocket deer
red brocket deer.............55
red deer . . 10, 22-25, 29, 37, 38, 40, 41,
46, 48-50, 53, 54, 56-58, 67-
69, 71, 73-81, 84, 85, 89, 92-
94, 97, 104, 109, 110, 114,
120, 122, 125, 132, 133, 136,
138-140, 144, 145, 151, 152,
155-158, 162-169, 173-178,
180, 182, 197, 210, 212-214,
218, 220, 222, 223, 230, 232-
234, 237-241, 248, 255
Reeveโs muntjac ................91
regeneration . 18, 42, 50, 160, 161, 192,
195, 200, 210, 220
reintroduction ......23, 106, 128, 253
reproduction ..... 16, 31, 49, 56, 109,
112-114, 118, 138, 181, 183,
184, 194, 195, 252
contraception ..............114
non-seasonal reproduction .....16
reproductive performance . . 80, 183
reproductive period ..........45
seasonal reproduction.........16
reproductive capacity ...........138
reproductive lifespan ........138, 139
reproductive success 43, 115-117, 119,
128
resource defense ...............115
re-epithelialisation..............161
roe deer . 16, 22, 29, 50, 53, 54, 57, 58,
70, 74, 88, 136, 140, 154-156,
165, 178, 179, 204, 220
ruminal trematode ...............79
Paramphistomum cervi ........79
ruminants . . 10, 12, 39, 41, 81, 91, 179,
180, 183
rut . 16, 49, 56, 80, 117, 119, 125, 137,
138, 174-178, 213
sanitary monitoring ..............81
Scotland .......25, 84, 151, 213, 228
seasonality .............73, 219, 244
seed dispersal .................209
semen ....... 109, 114, 122, 187-189
sex . . 24, 25, 27, 45, 46, 48, 57, 59, 60,
69, 84, 85, 92, 93, 104, 119,
123, 124, 129, 133, 136, 138,
139, 154, 157, 168, 175, 223,
274 R
225, 226, 255
sex differences ............. 46, 123
sex-specific lifetime strategies
.................... 139
sex ratio ........ 24, 25, 57, 119, 157
population sex ratio .......... 24
sexes . . . 8, 17, 27, 45, 47, 48, 56, 114-
116, 130, 139, 165, 178, 219,
225, 239, 252
Sexual choice ................. 117
sexual dimorphism ............. 114
sexual segregation .... 17, 45, 50, 130
sexual selection ............... 115
sheep . . 5, 9, 11, 35, 36, 67, 92, 94, 97,
179, 180, 185
signposting ................... 116
skeleton development............ 28
Slovak Republic
Slovakia............... 78, 222
snow track density index ..... 24, 144
snow tracking .................. 58
Solenopotes ................... 82
Solenopotes burmeisteri ...... 82
South America
Argentina....85, 87, 96, 106, 107,
174, 181, 183-185, 216, 253
Bolivia................... 217
Brazil.....5, 56, 87, 96, 172, 211,
239, 254
Chile . 16, 106, 107, 181-183, 185,
252, 253
Panama ............... 55, 254
sperm
sperm concentration......... 114
sperm motility .............. 47
spermatozoa . . 114, 120, 122, 188, 189
sperms
cryopreservation of spermatozoa
.................... 122
spongiform encephalopathy . . 6, 8, 12,
15
spotted deer .................. 102
stable isotopes ................ 244
stag . . 36, 101, 104, 116, 152, 164, 168,
169, 176, 187, 188, 211, 222,
229, 241
stem cells ............. 18, 160-162
strain................ 9, 68, 74, 129
stress...... 68, 75, 109, 173, 234, 247
sucking .................. 133, 134
survival . . . 17, 23, 30, 31, 55, 106, 116,
128, 135, 152, 183-185, 252,
253
survival rate ................... 55
systematics ................ 85, 209
molecular systematics ........ 85
taxonomy ...................... 2
teeth ....... 59, 74, 75, 138, 139, 244
fluorotic teeth .............. 74
premolars.................. 59
telemetacarpalian ............... 85
telemetry....8, 44, 106, 107, 119, 130,
138, 140, 156, 183, 185, 192,
193, 204, 222, 249
radiotelemetry . 17, 22, 23, 48, 192,
198
temperature . . . 3, 48, 73, 125, 140, 148,
150, 187, 188, 196, 235, 238,
239, 244, 246
territory............... 57, 176, 177
testes........................ 187
theory ............ 17, 130, 137, 195
therapy .................... 76, 78
anthelmintic treatment ..... 72, 80
antitrematod treatment ........ 80
copper oxide wire particles .... 76
thermal infrared imaging
thermal infrared imager ...... 146
tooth wear.......... 59, 60, 138, 139
toxic metals .................. 240
cadmium ............. 240, 241
lead ...... 36, 111, 240, 241, 247
mercury .............. 240, 241
toxicity ...................... 240
Tragulidae .................... 91
Tragulus .................. 91
Tragulus
Tragulus javanicus .......... 91
trampling ..................... 50
trees ....36, 41, 42, 50, 153, 175, 181,
200, 209, 244, 245, 255
trematodes
liver fluke .............. 78, 79
ruminal trematode ........ 79, 80
R 275
trophy ........25, 144, 152, 158, 197
ungulate . . 41, 102, 115, 116, 131, 132,
153, 196, 220, 226, 252
urine ...................33, 70, 71
uterine .......................124
vaccination ........... 62, 68-70, 74
Variability Size Index
V.S.I. ....................248
Variability Size Index ........248
vegetation . 30, 34, 40, 42, 44, 48, 107,
110, 153, 154, 179, 194, 196-
200, 213, 225, 244, 247
velvet.... 94, 116, 164, 165, 168, 187,
189
vocalization...........114, 115, 175
hoarse vocalization..........114
roaring ...............125, 176
Volkmannโs canals .............168
wapiti . 29, 46, 64, 69, 71, 97, 108, 109
weaning . 58, 59, 80, 94, 118, 131, 133,
140, 167, 229
weaning date................80
weather ........................3
welfare .......30, 58, 62, 63, 66, 127
wild animals ...........5, 6, 15, 122
wild boar.............5, 29, 102
wildlife . . 2, 4-8, 10, 11, 13-15, 22, 24,
28, 29, 31, 36, 37, 39-42, 44,
48, 49, 53, 54, 65-67, 78, 100-
106, 108, 111, 114, 115, 118,
119, 121, 136, 144, 146, 150-
152, 173, 185, 186, 192-194,
204, 208, 210-212, 218
wildlife warning reflectors . . . 193, 194
wood ........................111
zooarcheology.................243
zoogeography ...................2
6
t
h
I
N
T
E
R
N
A
T
I
O
N
A
L
D
E
E
R
B
I
O
L
O
G
Y
C
O
N
G
R
E
S
S
P
R
A
H
A
,
C
Z
E
C
H
R
E
P
U
B
L
I
C
6
t
h
I
N
T
E
R
N
A
T
I
O
N
A
L
D
E
E
R
B
I
O
L
O
G
Y
C
O
N
G
R
E
S
S
P
R
A
H
A
,
C
Z
E
C
H
R
E
P
U
B
L
I
C
2006