Le système d'appariement de Cerfs Sika (Cervus nippon) introduits dans la forêt de Manetin (220 km2), République tchèque, a été étudié de 1989 à 1993. Le Sika était strictement nocturne en raison des pressions de chasse et des

        dérangements continus par l'homme dans la journée. Les activités de rut ont été suivies par triangulation des cris des mâles en divers sites, ainsi que par comptage au phare des cerfs dans les zones ouvertes. Durant toutes les saisons, les activités vocales ne furent pas réparties

        uniformément sur la zone d'étude, étant généralement concentrées dans des champs proches de la forêt. En 1989, un lek fut constitué. Par la suite, il éclata en arènes dispersées. Ces arènes furent occupées par intermittence et étaient localisées tant dans les pâturages ouverts qu'en forêt

        mature et à l'interface de ces deux types d'habitat

BARTOŠ, Ludek

HERRMANN, Hubert

LOSOS, Sobeslav

MIKEŠ, Jaroslav

ŠILER, Jiri

I-Revues

VARIATION OF MATING SYSTEMS OF INTRODUCED SIKA DEER 
Ludek BARTOS* ,  Hubert HERRMANN*, Jin SILER* , Sobeslav Losos*,
& Jaroslav MIIŒS** 
Sika deer (Cervus nippon) exhibit an unusual degree of variability in  their 
mating system. Direct harem defence has been recorded in New Zealand (Kiddie, 
1 962) and England (Putman & Mann, 1 990). While, resource-based territories 
have been reported for native sika in Russia (Yevtushevskiy, 1 974) and Japan 
(Miura, 1 984 ), and for introduced populations in En gland (Horwood & Masters, 
1 970) and Ireland (Gradl-Grams, 1 982) . In the Czech Republic, we observed lek 
territoriality in Sika in the Manetfn Forest, Western Bohemia (Bartas et al. , ( 1 99 1 ) . 
This elicited our attention to this population for a further four seasons (see also 
Balrnford et al. , 1 993) .  The only other deer species known to exhibit a comparable 
ftexibility of mating system appears to be the Fallow deer (Dama dama, Langbein 
& Thirgood, 1 989), and possibly the Red deer (Cervus elaphus, Carranza et al. , 
1 990; Carranza, 1 992; Pereladova, pers. comm.).  
This paper analyses mating systems according to classification by Langbein 
& Thirgood ( 1 989) and summarizes records of Sika rutting behaviour at Manetfn 
over five seasons from 1 989 to 1 993 . To make it complete, we have also used 
ex tracts from two previously published reports (Bartas et al. , 1 99 1 ;  Balmford et 
al. , 1 993) .  
METHODS 
Sika were studied from 1 989 to 1 993 in a ca 20 km2 section of the 220 km2 
Manetfn Forest. Vegetation consisted of a mixture of open pasture (43 %) and 
commercial forests (spruce 53 %,  pine 40 %, and other tree species 7 %). Sika at 
Manetfn are strictly nocturnal as a result of hunting pressure, disturbance by 
tourists, and people gathering mushrooms during the daylight hours . We, therefore, 
monitored rutting activity by recording the direction of male calling from twelve 
different sites (Bartas et al. , 1 99 1 ;  Balmford et al. , 1993) .  These sites were 
distributed attempting to caver the whole area. Male calling was beard exclusively 
after sun set. Observations in 1 990 (Balmford et al. , 1993) demonstrated, th at 
* Research Institute of Animal Production, CZ- 1 04 00 Praha 10-Uhi'ineves, Czech Republic, 
e-mail: bartos@ novell .vuzv.cz or bartos @ vuzv.cz. 
** Forestry Company Pfestice, CZ-334 01 Pi'estice, Czech Republic. 
Rev. Eco[. (Terre Vie), vol. 53 ,  1 998.  
- 337 -
calling peaked at about 2200 hours and declining remarkably after midnight. We 
listened from four or more of these sites between 1 900 and 2200 hours for one day 
( 1 99 1 ), two days ( 1 989, 1992), or three days ( 1 993) at the usual peak of the rut 
( 14th to 1 6th October). Local foresters suggested the date of observations as they 
were able to estimate the approach of the peak of the rut. In 1 990 recording lasted 
three weeks (Balmford et al. , 1 993), however, only data for the analogous three 
days of 1 990 are used in this study. ln each year, we were able to cover most of 
the study area. We recorded the time and direction of each male calling bout, each 
bout consisting of one to four calls (Miura, 1 984). When calling bouts were beard 
simultaneously, from more than one monitoring site, the locations of calling males 
were determined by triangulation. Such locations are referred to as ' sites ' . If 
calling bouts were only beard from a single monitoring site, they were omitted 
from this study. From 199 1  onwards we counted Sika in open areas using a spot 
light from a car, both before (approximately between 1 800 and 1 900 hours) and 
after (approximately between 2200 and 2300 hours) recording sessions. However, 
it was not usually possible to visually determine the number of males at each site. 
We classified the observed activities according to definitions given by 
Langbein & Thirgood ( 1 989) . Territorial activities were regarded in the three 
following categories : 
'Stand' A single mature male defends an exclusive reproductive 
territory. Stands may vary greatly in habitat type and in size 
and may be clumped or widely dispersed. 
'Temporary stand' - A similar strategy to the above but one in which the stand 
'Lek' 
is only defended for a relatively small proportion of the day. 
range in composition from 5 to 25 mature males, charac­
terized by a reduction of individual terri tory size to as little 
as 5 x 5 m. 
According to these definitions we have forrnulated presumptions for indi­
vidual categories (Table 1) and applied them to the data collected. With a limited 
possibility to observe the mating activity visually, temporary stands appear 
difficult to distinguish from non-territorial strategies . 
TABLE 1 
Presumptions for individual categories of territorial strategies. 
Activity 
Calling sites widely dispersed over the area 
Calling activity reduced to a small propor-
tion of the day 
Females in proximity to calling males 
Repeated calling activity in the same site in 
successive da ys 
Local calling frequency higher th an in 
other sites 
Reduction of the distance between calling 
males to only a few metres 
338 
Stand 
Y es/No 
No 
Y es/No 
Y es 
No 
No 
Temporary stand 
or non-territorial 
strategies 
No 
Y es 
Y es 
Y es/No 
No 
No 
Lek 
No 
Y es 
Y es 
Y es/No 
Y es 
Y es 
If not stated otherwise, the data were analysed according to the General 
Linear Models Procedure (GLM) for Unbalanced ANOVA (SAS). The classes used 
were 'Caliing frequency of the site ' ( 'A site with the highest caliing frequency' and 
'Other sites ' )  and 'Season' ( 1 989 to 1 993). Least-squares means (LSMEAN) were 
computed for each class and differences between classes were tested using a t-test. 
RESULTS 
DISTRIBUTION OF CALLING SITES 
The distribution of locations for caliing males, over the years studied, is 
shown in Fig. 1 .  In each monitored season, caliing activity was unequaliy 
distributed over the area, usualiy concentrated in fields near the forest surrounding 
the smali village of Lipf. The maps in Fig. 1 show ca 20 % of the area covered by 
the observers ; there were no records of Sika caliing in the rest of the area (mostly 
forested habitat). 
PROXIMITY OF FEMALES TO CALLING MALES 
The results of observations of deer seen during the spot light searches are 
shown in Fig. 2 .  We could only compare the locations of caliing activity with 
visual records of deer numbers when the caliing locations were outside the forest 
and hence visible from a car. In ali 10 instances where such a comparison was 
possible, caliing activity coincided with the presence of deer of both sexes (Sign 
test, P < 0.00 1 ) .  No significant relationship was found between caliing frequency 
and the number of deer present at these sites (Spearman rank correlation rs = 0.4 1 ,  
n = 1 0, NS). 
REPETITION OF CALLING IN SAME SITES ACROSS SEASONS AND IN SUCCESSIVE 
DAYS 
When possible we compared calling on successive nights to establish whether 
sites were consistently occupied. Calls were heard from the same site on 
consecutive nights in 67 % ( 1 989, n = 3 sites), 47 % ( 1990, n = 1 7  sites), 25 % 
( 1 992, n = 8 sites), and 64 % ( 1 993,  n = 14  sites) of the cases. Across ali seasons, 
only 50 % of sites were occupied on successive nights . This suggests that the sites 
do not necessarily represent permanent territories. 
DIFFERENCES IN CALLING FREQUENCIES BETWEEN SITES WITHIN AND ACROSS 
SEASONS 
We calculated frequency of Sika male caliing bouts, by hour, for ali sites 
daily. In ali seasons,  certain sites showed a higher calling frequency than the other 
sites. Proportions of ali caliing bouts within the session at these most active sites 
were 92. 1 6 % ( 1 989), 77. 1 7 % ( 1 990), 50.67 % ( 1 99 1 ), 56 .88 % ( 1 992), and 
- 339 -
0.5 1 km 
Figure 1 .  - Locations of calling sika males 
between 1989 and 1993. 
Black spots : indicate at )east one calling bout; 
White arrow: the lek; Black arrow: the site with 
the most calling bouts ; Shaded part of the map: 
forests; White part of the map: fields . 
340 
O.S 1 km 
Figure 2. - Distribution of deer observed by 
spot light counts in 1991  to 1 993. 
Black spots : indicate at !east one animal seen 
during ali observations of the season; Black 
arrow: the site with the highest proportion of 
calling bouts per season; Areas: marked with 
dotted line show listening sites derived from 
Fig. 1 ;  Shaded part of the map: forests; White 
part of the map: fields. 
58 .75 % ( 1 993) .  For statistical analysis, data from ali seasons were combined and 
subjected to GLM. Calculation was carried out in two steps .  
First, we compared seasons. The GLM mode! containing the only class 
'Season' showed significant variation in calling frequency between seasons 
(inserted part in Fig .  3, F5 •4 1  = 4.83 ,  P < 0.0 1 ) . Calling frequency in 1 989 was
higher than in 1 990 (P < 0.05), 1 992 (P = 0.07), and 1993 (P = 0.06), but not in 
1 99 1  (P = 0. 1 67 1 ) . The calling frequencies in ali other seasons did not significantly 
differ from each other. 
Second, we compared hourly calling frequencies between the sites with the 
highest calling activity and the other sites (Fig. 3) .  For this, the class 'Calling 
frequency of the site ' was nested within 'Season' . There was a significant variance 
between sites and seasons (F9•4 1  = 1 5 .40, P < 0.00 1 ) .  Only in 1 989 did the site with
the highest calling frequency produce significantly more calling than the other sites 
(P < 0 .005) .  
34 1 
1... 70 :::J 
0 
� 60 >. � 50 
lj' 
c: 40 Q) 
:::J 
� 30
-
Cl 20 c: 
ca 
(.) 1 0
0 
n= 2 1 5
1 990 
3 / 9  
1 990 
4 0  ,.....-----------, 
30 
20 
1 0  
0 
n=7 n= 1 2  n=3 n=9 n=7 
1 989 1 990  1 99 1  1992 1993 
1 / 2 2 / 7
1 99 1  1 992 
Season 
3 / 8
1 993 
D Sites with the highest cal l ing frequency
• Other sites
Figure 3 . - The mean houri y frequency (LSMEAN ± S .E.)  of Sika male calling bouts per site per day, 
and sites with the highest proportion of the records across seasons. Inserted part shows calling 
frequency (LSMEAN ± S .E.) between seasons. Black bars : Calling bouts per site per day; White bars: 
Calling bouts per site with the highest activity per day. 
It is interesting to note that since 1 99 1 ,  calling was frequently beard in village 
gardens within tens of metres of the bouses. In 1 99 1  this site bad even the highest 
calling activity (Fig. 1 ,  middle left) . 
THE DISTANCE BETWEEN CALLING MALES 
In 1 989 the shortest distance between calling males was approximately 40 rn 
on a putative lek, when up to 7 males were rutting on a spot of about 300 rn in 
diameter (Bartos et al. , 1 99 1 ) .  In ali other seasons the inter-male distance did not 
fall below several hundred metres .  
During daylight we exarnined calling sites in order to quantify pawing or 
scraping activity, as a possible sign of territorial behaviour. Although in 1 989, the 
lek did contain clusters of scrapes (ibid.),  in other seasons we found as many 
scrapes at other sites as at calling sites. 
342 -
DISCUSSION 
In the studied population of Sika deer, the mating system bas evidently 
changed across seasons. A typical resource-based territorial system, as described in 
most native or introduced Sika populations (Horwood & Masters, 1 970; Yevtush­
evskiy, 1 974; Gradl-Grams, 1 982; Miura, 1 984), appears unlikely in the Manetin 
Forest. Although conditions for true lekking (Table I) were fulfilled only for 
observation in 1 989 (Bartos et al. , 199 1 ), from 1 990 to 1 993 calling males were 
also aggregated, and in each mating season the distance between extreme calling 
positions did not usually exceed 2 km. In contrast to true lekking behaviour, 
however, rutting stands in these later years were usually located on or close to 
resources used by females. 
Where we could monitor deer both visually and acoustically, we usually 
found that calling sites were located in areas with concentrations of deer (Fig. 2). 
These sites were not necessarily in the same place on consecutive nights . Instead, 
it seemed that calling males moved into areas where the hinds bad relocated. This 
may explain why we failed to find any consistent relationship between scrape 
location and calling activity. In sorne other species scraping is facultative and not 
necessarily linked to territory marking (Hirth, 1 977). In White-tailed deer, scraping 
serves as a means of communication between bucks and does during the breeding 
season (Kile & Marchinton, 1 977;  Miller et al. , 1987 ; Ozoga, 1 989), with most 
scrapes being made by mature, dominant, males to intimidate rival males and to 
attract breeding females (Marchinton & Hirth, 1984). Marchinton & Atkeson 
( 1 985) suggested that territoriality in White-tailed deer is facultative, and depends 
on resource distribution and population demography. This facultative interpreta­
tion of territoriality may also explain its absence in Sika deer in Manetfn, where 
permanent harassment by humans during daylight prevents the males from 
establishing themselves in fixed locations (Bartos et al. , 1 99 1 ) .  
We conclude that Sika males i n  Manetin occupy temporary rutting stands 
located in open pasture, mature forest, and in the boundaries between these two 
habitats. Males also called extensively from within the village gardens. These 
stands do not seem to be permanent resource-based territories held throughout the 
mating season (reported Yevtushevskiy, 1 974 and Miura, 1 984). Instead, the stands 
are clustered, temporarily, in areas where resources used by females are particu­
larly concentrated, or on paths used by females commuting between resources. 
This gives the deer advantages similar to those arising through lekking; females 
benefit from increased opportunity to choose a mating partner quickly, they are 
able to compare males closely and also have the possibility to monitor choices of 
other females (Bradbury & Gibson, 1 983 ; Clutton-Brock et al. , 1988; Clutton­
Brock et al. , 1 989). Nevertheless, it is apparent that because the males are more 
dispersed, the opportunities for females to assess and compare them are less than 
on a lek. True lekking in Sika seems to occur only in high density populations 
(Bartos et al. , 1 99 1 ;  Balmford et al. , 1 993).  Sika thus prove to be successful 
opportunists that can adapt ftexibly to changing environments. 
ACKNOWLEDGEMENTS 
We wish to thank Andrew Balmford, R. Larry Marchinton, P. Duncan, and J.M. Gaillard for 
comrnents on the manuscript, Jon Benge and Jasmin Barwig for their help with preparation of the 
English version of the manuscript, and to Petr Janovsky, Josef Mika, Jitka Lancingerovâ and 
Dominika Vaiinkovâ for assistance in field observations. 
- 343 -
SUMMARY 
The mating system of ferai Sika deer (Cervus nippon) in the Manetfn Forest 
(220 km2), Czech Republic, was studied from 1 989 to 1 993 .  The Sika were strictly
nocturnal due to hunting pressure and continuous human harassment during 
daylight. We therefore monitored rutting activity by triangulating male Sika calls 
recorded from different sites, and also by counting Sika in open areas using a spot 
light. In all seasons, calling activity was unequally distributed over the area, 
usually concentrated in fields close to the forest. In 1 989 the Sika established a lek. 
Later, the mating system changed to dispersed rutting stands. These rutting stands 
were occupied intermittently and were located in open pasture, mature forest, and 
in the boundaries between these two habitat types. 
RÉSUMÉ 
Le système d' appariement de Cerfs Sika (Cervus nippon) introduits dans la 
forêt de Manetin (220 km2), République tchèque, a été étudié de 1 989 à 1 993 .  Le 
Sika était strictement nocturne en raison des pressions de chasse et des dérange­
ments continus par l 'homme dans la journée. Les activités de rut ont été suivies par 
triangulation des cris des mâles en divers sites,  ainsi que par comptage au phare 
des cerfs dans les zones ouvertes. Durant toutes les saisons, les activités vocales 
ne furent pas réparties uniformément sur la zone d'étude, étant généralement 
concentrées dans des champs proches de la forêt. En 1 989, un lek fut constitué. Par 
la suite, il éclata en arènes dispersées .  Ces arènes furent occupées par intermittence 
et étaient localisées tant dans les pâturages ouverts qu' en forêt mature et à 
l ' interface de ces deux types d' habitat. 
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Variation of mating systems of introduced sika deer

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Variation of mating systems of introduced sika deer

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