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How does parental role influence the activity and movements of breeding wolves?

DOI:10.1007/s10164-008-0106-z
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Hiroshi Tsunoda
Hiroshi Tsunoda
Hiroshi Tsunoda
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Roman Gula at Polish Academy of Sciences
Roman Gula
Jörn Theuerkauf at Polish Academy of Sciences
Jörn Theuerkauf
Sophie Rouys
Sophie Rouys
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We studied the activity and movements of a male and a female wolf during breeding in the Bieszczady Mountains, Poland. The female was less active and mobile and on average closer to the den than the male. The male was less active and mobile when the pack size was seven compared to a year when the pack consisted only of the breeding pair. We conclude that the roles played by breeding males and females rearing pups influence their activity, and that breeding males in larger packs move less during the nursing period because of help by other pack members.
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SHORT COMMUNICATION
How does parental role influence the activity and movements
of breeding wolves?
Hiroshi Tsunoda Æ Roman Gula Æ Jo
¨
rn Theuerkauf Æ
Sophie Rouys Æ Stephan Radler Æ Bartosz Pirga Æ
Julia Eggermann Æ Barbara Brzezowska
Received: 15 January 2008 / Accepted: 3 June 2008 / Published online: 10 July 2008
Ó Japan Ethological Society and Springer 2008
Abstract We studied the activity and movements of a
male and a female wolf during breeding in the Bieszczady
Mountains, Poland. The female was less active and mobile
and on average closer to the den than the male. The male
was less active and mobile when the pack size was seven
compared to a year when the pack consisted only of the
breeding pair. We conclude that the roles played by
breeding males and females rearing pups influence their
activity, and that breeding males in larger packs move less
during the nursing period because of help by other pack
members.
Keywords Bieszczady Mountains Canis lupus
Feeding helpers Poland Reproduction
Introduction
Little is known about the social behavior of free-ranging
wolves (Canis lupus), as these animals are difficult to
observe and most behavioral studies therefore deal with
captive packs (e.g., Ziemen 1982; Asa et al. 1990; Bernal
and Packard 1997). With a few exceptions (e.g., Mech
H. Tsunoda
Wildlife Conservation Laboratory, Graduate School of
Agriculture, Tokyo University of Agriculture and Technology,
Saiwai-cho 3-5-8, Fuchu-shi, Tokyo 183-8509, Japan
Present Address:
H. Tsunoda
Laboratory of Water Resource Planning,
United Graduate School of Agricultural Sciences,
Tokyo University of Agriculture and Technology,
Saiwai-cho 3-5-8, Fuchu-shi, Tokyo 183-8509, Japan
e-mail: dayan_and_jitan@hotmail.com
R. Gula J. Theuerkauf B. Pirga
Museum and Institute of Zoology, Polish Academy of Sciences,
Wilcza 64, 00-679 Warsaw, Poland
e-mail: jtheuer@miiz.waw.pl
B. Pirga
e-mail: wrzosowe_wzgorze@poczta.onet.pl
S. Rouys
Conservation Research New Caledonia, BP 2549,
Noumea 98846, New Caledonia
e-mail: rouys@ifrance.com
S. Radler
University of Applied Forest Sciences Rottenburg,
Schadenweilerhof, 72108 Rottenburg, Germany
e-mail: stephan.radler@fh-rottenburg.de
J. Eggermann
Faculty of Biology and Biotechnology,
Ruhr University Bochum, Universita
¨
tsstr. 150,
44801 Bochum, Germany
e-mail: julia.eggermann@web.de
B. Brzezowska
Department of Animal Ecology,
Institute of Environmental Sciences,
Jagiellonian University, Gronostajowa 7,
30-387 Krakow, Poland
e-mail: bulbulek@wp.pl
R. Gula (&)
45 rue Maurice Herzog, Noumea 98800,
New Caledonia
e-mail: rgula@miiz.waw.pl
123
J Ethol (2009) 27:185–189
DOI 10.1007/s10164-008-0106-z
1999), studies of wild wolves are accordingly based on
indirect methods, such as telemetry. Behavioral information
is therefore often gathered as a byproduct rather than an
answer to a primary question. The social and physiological
status of free-ranging wolves has an impact on the activity
and movements of individuals (Harrington and Mech 1982;
Asa et al. 1990; Ballard et al. 1991), which can vary with
age and sex (Je˛drzejewski et al. 2001; Schmidt et al. 2008),
dominance (Mech 1999) and reproductive status (Harring-
ton and Mech 1982; Ballard et al. 1991; Vila
`
et al. 1995;
Je˛drzejewski et al. 2001; Theuerkauf et al. 2003a). These
intrinsic factors are mitigated by environmental factors
which also shape the activity and movements of wolves.
Documented external factors that influence wolf activity
and movements are prey density (Messier 1985), prey
migration (Walton et al. 2001), prey activity (Theuerkauf
et al. 2003a), the activity of neighboring packs (Mech and
Harper 2002), and human activity (Vila
`
et al. 1995; Ciucci
et al. 1997; Theuerkauf et al. 2003a, b; Kaartinen et al.
2005; Kusak et al. 2005; Theuerkauf et al. 2007).
During breeding, wolves gravitate around den and ren-
dezvous sites (summarized in Mech and Boitani 2003).
Besides the breeding male, nonreproductive adult pack
members also supply the nursing mother and pups with
meat (Harrington et al. 1983; Mech et al. 1999). This al-
loparental care has an impact on the pups’ survival
(Harrington et al. 1983) and should also influence the
activity budget of the breeding male. We expected that in
the absence of help from other nonbreeding adults, the
breeding male, who would then be the sole food supplier,
should increase his activity and movements. On the other
hand, in a larger pack, the breeding male could remain
longer with his mate and pups to protect them. To verify
this hypothesis, we compared the activity and movements
of a breeding female and of a breeding male in a year when
he was alone with his mate with those in a year when there
were seven pack members.
Methods
The study area is situated in the Bieszczady Mountains,
southeastern Poland (49°19
0
–49°50
0
N, 22°15
0
–22°45
0
E) and
covers approximately 1,000 km
2
. Elevations range from
300 to 900 m a.s.l. The mean annual temperature is 5.5 °C
and the annual precipitation ranges from 800 to 1,200 mm.
Forest, mostly beech (Fagus sylvatica), fir (Abies alba),
spruce (Picea abies) and grey alder (Alnus incana), covers
62% of the area. Wolf packs consist of 2–7 individuals and
occupy home ranges of 88–229 km
2
(Gula 2008).
We radiotracked two breeding wolves (one male and
one female) from different packs. Wolves were fitted with
VHF radiocollars without activity sensors (Telonics Inc.,
Mesa, AZ, USA). The female was caught in March 2002.
From the beginning of May 2004 and for the following
40 days she remained mostly in one place, suggesting that
she was rearing pups. In July, we investigated that spot and
found her den. In August and September, howling revealed
that her pack also included pups. We therefore considered
that the female had raised young in 2004.
The second wolf was a three-year-old dispersing male
when caught in spring 2003. He was alone when snow-
tracked in December 2003, but from January 2004 onward
he was accompanied by another wolf, apparently a female,
as we found signs of mating in the snow. This pair must
have bred in 2004, as we heard them howl with pups. The
following summer, based on radio-locations of the male,
we found their den. In the summers of 2005 and 2006 we
also heard pups howling with several adults, so the pack
must have bred in those years too. By snow-tracking, we
estimated that the pack consisted of five individuals in the
winter of 2004/2005 and seven in the winter of 2005/2006.
We estimated the parturition date as the first time that we
located wolves at the spot where we later found a den (in
our study area during the first ten days of May).
In this paper, we used radiotracking data for the female
collected in 2004 and for the male collected in 2004 and
2006. Our data cover the period from mid May to early
September, during which each wolf was followed every
second week over a whole 24-h period. During these 24-h
sessions, we located wolves by ground triangulation and
recorded wolf locations and activity (based on changes in
signal strength) every 15 min. To estimate the proportion
of time active, we assigned a value of 1 when the wolves
were active and their locations changed; a value of 0 when
the wolves were not active and their locations did not
change; and a value of 0.5 when the wolves were either
active or their locations changed (as described in The-
uerkauf and Je˛drzejewski 2002). We estimated the distance
to the den as the straight-line distance from a wolf location
to the den site. The location error was about 250 m (The-
uerkauf et al. 2007). Because we could not find the den of
the male in 2006, we estimated the location of the den site
as the center of the activity of the male in May and June of
2006 (geometric center of the 25% kernel density distri-
bution of locations). Later, we heard the pack howl with
pups at this site. We estimated the daily range of each
radiotracking session as minimum convex polygons
(MCPs) of the 96 locations taken during the 24 h.
Results
As the breeding season progressed, both wolves increased
the distance they traveled from the den (Fig. 1). The time
active, the distance traveled and the daily range of the
186 J Ethol (2009) 27:185–189
123
female increased gradually, whereas den attendance and
the time she spent at the den steadily decreased. In 2004,
when the breeding pair was alone, the male’s activity, the
distance traveled and the daily range were highest just after
the pups’ birth. These values constantly decreased until
about three months after birth, when they reached about the
same levels as those of the female (Fig. 1). In 2004, the
breeding male was mostly away from the den and his
activity and movements were accordingly greater than
those of the female. In 2006, however, when the male’s
Fig. 1 Seasonal changes in mean distance to the den, time spent at
the den, den attendance, distance traveled, time spent active and daily
range of a female (n = 6 days in 2004) and male wolf (n = 5 in 2004
and n = 5 in 2006) during two breeding seasons in the Bieszczady
Mountains, SE Poland
J Ethol (2009) 27:185–189 187
123
pack consisted of seven wolves, his activity pattern and
movements were reduced to levels that were similar to
those of the female (Table 1).
The activity and distance traveled by the female were
higher in the day and at twilight than at night (Table 1),
and most differences between years or individuals occurred
at night. In 2004, the male’s nightly movements were more
than twice as high as they were in 2006. The female, on the
other hand, moved very little at night (Fig. 2). These trends
were also reflected in the time spent at the den. In 2004, the
male spent virtually no time at the den, especially in the
twilight hours. In 2006, however, the male spent compa-
rable amounts of time at the den to the female in 2004.
Accordingly, the male’s daily range was much larger in
2004 (11.3 ± 3.6 km
2
95% confidence interval) than in
2006 (3.2 ± 3.1 km
2
). The female’s range was the smallest
of all (2.5 ± 1.7 km
2
).
Discussion
Although other factors, such as prey availability, might have
influenced the activity of the wolves, we think that the
differences in male and female behavior were a result of their
different roles in the biparental care of pups. Mothers nurse
the pups, keep them warm and keep the den clean and dry
(summarized in Packard 2003). In our study, the female left
the den mostly in the day, which would correlate with her
having to remain with the pups to keep them warm during
cooler periods (twilight and night). Breeding males, on the
other hand, attend the den site less frequently than the adult
females (Harrington and Mech 1982; Ballard et al. 1991;
Potvin et al. 2004), but contribute by feeding the female and
therefore spend most of the time hunting away from the den
(Mech 1999). Wolves in our study area probably hunt mostly
at dawn (Eggermann et al. 2008), which would explain why
this was the time when the breeding male was almost never at
the den. Previous studies report that adult females with pups
were less active during the denning season (Ballard et al.
1991; Mech et al. 1995; Vila
`
et al. 1995;Je˛drzejewski et al.
2001; Theuerkauf et al. 2003a; Schmidt et al. 2008). The
activity of the female we monitored in this study was lower
than the mean activity of wolves in our study area (Egger-
mann et al. 2008), suggesting that she too reduced her
activity while she cared for the pups.
Despite tracking only one breeding male, to our
knowledge this is the only account of a breeding male’s
daily activity and movements during the denning season.
We observed a shift in his behavior after pack size
increased from two to seven individuals. In both years, the
breeding male’s activity and movements seem to have been
strongly linked to the necessity of supplying his mate, and
later pups, with food. His reduction in activity and move-
ments in 2006 support this hypothesis, since he was no
longer the sole food provider for his pack in that year.
About two months after their birth, although they are not
yet able to follow the adults on their extensive movements,
the pups are no longer dependent on their mother for milk
or warmth, and so they can be left alone at rendezvous
sites (summarized in Packard 2003). From this time on,
the breeding pair can resume their normal activity and
Table 1 Mean (with 95% confidence interval) time spent active, distance traveled and daily range of a female (n = 6 days in 2004) and a male
wolf (n = 5 in 2004 and n = 5 in 2006) during two breeding seasons in the Bieszczady Mountains, SE Poland
Sex, year Whole day (24 h) Daylight (13.5 h) Twilight (4 h) Night (6.5 h)
Time spent active (%) Male, 2004 44.2 ± 10.3 33.5 ± 8.3 51.9 ± 22.0 65.0 ± 30.6
Male, 2006 26.5 ± 8.2 20.3 ± 3.1 37.5 ± 11.6 30.4 ± 22.9
Female, 2004 23.1 ± 9.7 30.5 ± 17.8 30.7 ± 11.4 5.7 ± 3.9
Distance traveled (km/h) Male, 2004 0.67 ± 0.14 0.38 ± 0.25 0.77 ± 0.38 1.23 ± 0.79
Male, 2006 0.33 ± 0.14 0.14 ± 0.06 0.44 ± 0.35 0.53 ± 0.37
Female, 2004 0.30 ± 0.17 0.40 ± 0.30 0.49 ± 0.21 0.00 ± 0.00
Time spent at the den (h) Male, 2004 0.3 ± 0.5 0.1 ± 0.1 0.0 ± 0.0 0.3 ± 0.5
Male, 2006 7.4 ± 8.9 4.6 ± 5.5 1.2 ± 1.6 1.6 ± 2.3
Female, 2004 5.8 ± 6.0 2.7 ± 2.9 1.2 ± 1.4 1.9 ± 2.1
Fig. 2 Daily pattern of distance traveled by a female and a male wolf
during the 2004 and 2006 breeding seasons in the Bieszczady
Mountains, SE Poland
188 J Ethol (2009) 27:185–189
123
movements, which probably explains why, in 2004, the
activity and movements of the male and the female reached
the same levels about three months after birth. We there-
fore suggest that the activity patterns of male and female
breeding wolves are shaped by their differing roles during
the denning period and by the availability of feeding
helpers.
Acknowledgments This study was part of the Bieszczady Wolf
Project and was funded by the Polish National Committee for Sci-
entific Research (KBN 6P04F 006), and the Museum and Institute of
Zoology of the Polish Academy of Sciences. Fellowships were pro-
vided by the Japan Student Services Organization (to HT), the
German Donors’ Association for the Promotion of Sciences and
Humanities (to JT), and the German Academic Exchange Service (to
JE). We thank N. Maruyama and K. Perzanowski for their coopera-
tion and L. Aubry, M. Barreteau, M. Carruthers, M. Diemert, S.
Drevet, M. Januszczak, S. Kiener, K. Lahongre, M. Le Peutrec, K.
Mayer, N. Schmidt, M-C. Schultz, M. Skuban and W. Schwimmer for
their help during field work, as well as two anonymous reviewers for
useful comments.
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... Wolf activity is highly flexible and may be adapted to various environmental conditions with a peak of activity during summer, which corresponds to the nursing period (Fancy & Ballard 1995, Eggermann et al. 2008; however, breeding females decrease their mobility during summer (Eggermann et al. 2008). The reasons for lower mobility during summer are denning and nursing, when wolf activity concentrates on the den site and the breeding female does not move far from the den (Eggermann et al. 2008, Tsunoda et al. 2009). The intensity of territorial marking predicts wolf reproduction (Llaneza et al. 2014). ...
... We defined from a posteriori analysis the "nursing territory" as the territory comprised in a 4 km radius from the den (corresponding roughly to a 50 km 2 area). We chose this distance because a previous study by Tsunoda et al. (2009) showed that the mean travel distance from the den for radio-tracked wolves of both sexes during the nursing period rarely exceeded a total distance (round trip) of 8 km/night or day, with a mean daily range of 11.3 ± 3.6 km 2 centred around the den. Consequently, seven marking sites were included in the nursing territory. ...
... However, one must remember that young wolves did not help the Neowise wolf-pack to provide food to the female as other wolf packs do; hence, the defecation patterns may be different if one or more young wolves from a previous reproduction help the breeding couple parenting the pups. This hypothesis seems likely, as a previous study from Tsunoda et al. (2009) showed that a radio-collared male (of a breeding pair raising the pups without aid, as in our study) was mostly away from the den and his activity and movements were accordingly greater than those of the female during the nursing season. In the following years, when the male's pack consisted of seven wolves, his activity pattern and movements matched those of the female. ...
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... However, unlike most other large carnivores, wolves have a complex social system, with a basic social unit (i.e., a pack) composed by a breeding pair and their offspring of one or more litters (Mech, 1999;Mech and Boitani et al., 2003). The nonbreeders have an important role in this social system, contributing as helpers for the protection and nursing of the mother and pups (Harrington et al., 1983;Mech, 1999;Clutton-Brock, 2002;Mech and Boitani et al., 2003;Tsunoda et al., 2009;Ruprecht et al., 2012;Ausband et al., 2016), which may greatly affect breeding success (Harrington et al., 1983;Sparkman et al., 2011) and the fitness of breeders (Mech, 1999;Tsunoda et al., 2009). Despite the paucity of comparative studies, this basic system seems to be considerably flexible, including for instance the possibility of packs incorporating unrelated individuals (Fritts and Mech, 1981;Van Ballenberghe, 1983;Peterson et al., 1984;Mech, 1987;Smith et al., 1997;Jimenez et al., 2017), the occurrence of simultaneous breeding by more than one female within the same pack (Smith et al., 1997;Mech and Boitani et al., 2003), or the wide variation in pup nurturing, and movement and activity patterns of breeders and non-breeders throughout the breeding season (Ballard et al., 1991;Harrington and Paquet, 1982;Ruprecht et al., 2012;Schmidt et al., 2008;Thurston, 2002;Tsunoda et al., 2009). ...
... However, unlike most other large carnivores, wolves have a complex social system, with a basic social unit (i.e., a pack) composed by a breeding pair and their offspring of one or more litters (Mech, 1999;Mech and Boitani et al., 2003). The nonbreeders have an important role in this social system, contributing as helpers for the protection and nursing of the mother and pups (Harrington et al., 1983;Mech, 1999;Clutton-Brock, 2002;Mech and Boitani et al., 2003;Tsunoda et al., 2009;Ruprecht et al., 2012;Ausband et al., 2016), which may greatly affect breeding success (Harrington et al., 1983;Sparkman et al., 2011) and the fitness of breeders (Mech, 1999;Tsunoda et al., 2009). Despite the paucity of comparative studies, this basic system seems to be considerably flexible, including for instance the possibility of packs incorporating unrelated individuals (Fritts and Mech, 1981;Van Ballenberghe, 1983;Peterson et al., 1984;Mech, 1987;Smith et al., 1997;Jimenez et al., 2017), the occurrence of simultaneous breeding by more than one female within the same pack (Smith et al., 1997;Mech and Boitani et al., 2003), or the wide variation in pup nurturing, and movement and activity patterns of breeders and non-breeders throughout the breeding season (Ballard et al., 1991;Harrington and Paquet, 1982;Ruprecht et al., 2012;Schmidt et al., 2008;Thurston, 2002;Tsunoda et al., 2009). ...
... The nonbreeders have an important role in this social system, contributing as helpers for the protection and nursing of the mother and pups (Harrington et al., 1983;Mech, 1999;Clutton-Brock, 2002;Mech and Boitani et al., 2003;Tsunoda et al., 2009;Ruprecht et al., 2012;Ausband et al., 2016), which may greatly affect breeding success (Harrington et al., 1983;Sparkman et al., 2011) and the fitness of breeders (Mech, 1999;Tsunoda et al., 2009). Despite the paucity of comparative studies, this basic system seems to be considerably flexible, including for instance the possibility of packs incorporating unrelated individuals (Fritts and Mech, 1981;Van Ballenberghe, 1983;Peterson et al., 1984;Mech, 1987;Smith et al., 1997;Jimenez et al., 2017), the occurrence of simultaneous breeding by more than one female within the same pack (Smith et al., 1997;Mech and Boitani et al., 2003), or the wide variation in pup nurturing, and movement and activity patterns of breeders and non-breeders throughout the breeding season (Ballard et al., 1991;Harrington and Paquet, 1982;Ruprecht et al., 2012;Schmidt et al., 2008;Thurston, 2002;Tsunoda et al., 2009). ...
Use of space and homesite attendance by Iberian wolves during the breeding season
Article
  • Mar 2018
The persistence of large carnivores in human-dominated landscapes is conditional on the preservation of adequate ecological conditions during the reproduction period, when they may be particularly susceptible to human disturbance. However, little is known about the breeding behaviour of large carnivores in these landscapes, though this would be important for conservation management. Here we describe the space use and homesite attendance patterns of wolves inhabiting humanised landscapes of north-western Portugal, based on GPS tracking of 11 individuals in 2008-2013. Parturitions (N=3) occurred in late May, with pups remaining at natal dens for 24-85 days, after which they were moved to a first and then sometimes to a second rendezvous site. Two of these movements were associated with human disturbance events. Breeding females (N=4) spent a large proportion of time with pups during pre-weaning (2 months after birth), leaving only at night during short periods; thereafter they progressively reduced time at homesites (i.e, natal dens and rendezvous sites), and increased both daily travel distances and daytime activity. Throughout the pup-rearing season, breeding females restricted their movements to within 􀀀2km of homesites. Non-breeding females (N=4) showed significantly lower attendance rates than breeding females, particularly during pre-weaning, and they had much less restricted movements throughout the breeding season, with activity occurring throughout the circadian cycle. Non-breeding males (N=3) showed attendance and space-use patterns intermediate between breeding and non-breeding females, with movements largely concentrated within the same areas used by breeding females. Our findings suggest that breeding wolves may be particularly susceptible to human activities occurring within 2km of homesites, although further research is needed to assess the ultimate effect of such disturbance on reproductive success and pup survival.
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... Studies of wolves have documented several factors that may influence their activity patterns. The time periods, when animals are active during the day and movements are influenced by the social and physiological status of wolves, can vary with age and sex (Jedrzejewski et al., 2001;Theuerkauf et al., 2003a;Eggermann et al., 2009), dominance (Mech, 1999), reproductive (Vilà et al., 1995;Theuerkauf et al., 2003a;Tsunoda et al., 2008) and social affiliation (Mancinelli et al., 2019;Yachmennikova & Poyarkov, 2011). Besides these intrinsic factors, there are environmental factors that also modify the activity and movements of wolves, including availability and activity of prey species (Messier, 1985(Messier, , 2011Theuerkauf, et al., 2003a), as well as weather conditions and human activity (Vilà et al., 1995;Ciucci et al., 1997;Theuerkauf et al., 2003aTheuerkauf et al., , 2007Kusak et al., 2005;Chavez & Gese, 2006;Kaartinen et al., 2015;. ...
... In our study, two breeding females did not reduce their overall activity and movement during the period of reproduction. However, F1 was less active at night and dawn, probably indicating that other wolves may have provided her with food (Theuerkauf et al., 2003a;Tsunoda et al., 2008). In our study, breeding females did not spend much of their denning time, compared to previous studies (Ballard et al., 1991;Vilà et al., 1995;Jedrzejewski et al., 2001;Schmidt et al., 2008;Theuerkauf et al., 2003a,b;Mancinelli, 2017). ...
Daily activity patterns of wolves in open habitats in the Dauria ecoregion, Russia
Article
Full-text available
  • Oct 2021
There are very little data about daily activity patterns of Canis lupus (hereinafter – wolf) living in open arid habitats with low human density in Dauria. Therefore we have studied the influence of human activity, reproduction and weather conditions on daily patterns and duration of the activity of 17 GPS-collared wolves in the Daursky State Nature Biosphere Reserve, Russia, from 2015 to 2020. GPS-collars were equipped with acceleration sensors. Wolves were active 44% (± 0.02 SE) of the day and traveled 1.21 km/h (± 0.10 SE) on average. The mean duration of subsequent activity periods was 7.36 h (± 1.5 SD). The duration of the subsequent, inactivity period was 10.07 h (± 4.2 SD). Travelling speed significantly increased when wolves made extraterritorial forays from their home range to territories of neighbouring packs. The highest activity index corresponds to long-distance dispersing wolves. Weather conditions and human activity did not significantly effect wolves daily activity patterns. Wolves were generally less active and mobile during the cold season. All wolves showed crepuscular movement peaks. Five of the wolves’ movement patterns switched to diurnal eight cases when they conducted an extraterritorial foray crossing territories of neighbouring packs. We conclude that wolves’ daily activity patterns were mainly shaped by a combination of several factors.
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... Instead of spending time in the den, 'breeding males' invest more time leading the foraging and food provisioning during the pup-raising season (Mech, 1999). During this time, 'breeding males' also move around the vast areas to maintain the territories (Alfredéen, 2006;Tsunoda et al., 2008). When males or other individuals are away from the den or pack, their long-distance vocalisation (howl) plays a crucial role in locating the pack and conveying alarm in extensive areas (Mazzini et al., 2013;Watson et al., 2018). ...
Understanding wolf howls and their application in individual identification and population estimation
Thesis
Full-text available
  • Nov 2022
The Indian wolf is a Schedule I species in the Wildlife Protection Act 1972. It is now considered an Evolutionary Significant Unit (A adaptive variation significantly important for conservation) (Hennelly et al., 2021). Since they survive predominantly in a human-dominated landscape (Habib et al., 2021; Habib & Kumar, 2007), they face immense survival threats due to habitat degradation and man-animal conflict (Agarwala et al., 2010). Their population status has remained unassessed over the years due to difficulties associated with the population estimation of this visually cryptic long-ranging species (Cozzi et al., 2021). A few studies have suggested that around 1000 to 2000 (Sillero-Zubiri et al., 2004) wolves are left in India, but those are rough estimates without statistical support. Therefore, a non-invasive statistical tool is required to estimate this visually cryptic species. Since the howling survey is considered the most efficient monitoring tool for this visually cryptic species (Harrington & Mech, 1982), my study aimed to standardise a statistical tool to estimate the population of Indian wolves based on their howl. I have started my work with a single point of reference on Indian wolf vocalisation – a comparative study of Indian wolf howls with a few other subspecies (Hennelly et al., 2017). I began the study with howling survey responses and opportunistic recordings from captive and nine free-ranging packs of Indian wolves. Different harmonic call types were characterised using an unsupervised statistical tool and defined to generate baseline information about the vocal characteristics of the Indian wolf. Through unsupervised clustering, I found four distinct vocalisations using 270 recorded calls (Average Silhouette width Si = 0.598), which include howls and howl-barks (N = 238), whimper (N = 2), social squeak (N = 28), and whine (N = 2). Indian wolf howls have an average mean fundamental frequency of 422 Hz (±126), similar to other wolf subspecies. The whimper showed the highest frequency modulation (37.296±4.601) and the highest mean fundamental frequency (1708±524 Hz) compared to other call types. Less information is available on the third vocalisation type, i.e. ‘Social squeak’ or ‘talking’ (Mean fundamental frequency = 461±83 Hz), which is highly variable (coefficient of frequency variation = 18.778±3.587). Lastly, I identified the whine, which had a mean fundamental frequency of 906Hz (±242) and was similar to the Italian wolf (979±109 Hz). The study highlighted how ‘social squeak’ can be misidentified with the howl. They can be differentiated through their frequency modulation and duration. Social squeaks (x̅ = 3.87s) are generally shorter than howl (x̅ = 5.214s). My study on the characterisation of the harmonic vocal repertoire provides a first step in understanding the function and contextual use of vocalisations in the Indian wolf. Studies over the years found that wolf howls contain individual-specific information (Fentress, 1967; Root-Gutteridge et al., 2014b, 2014a; Tooze et al., 1990). But identifying the unknown individual from their howls had remained challenging over the years, without which howl could not be used in Capture-Mark-Recapture studies (Marques et al., 2013; Stevenson et al., 2015). By understanding the importance of howl identification to an individual in population estimation, I trained a supervised model using known howls to identify howls to individuals. I verified the model with a set of unknown howls (unknown to the model). In this supervised classification, I achieved 97.9% accuracy in identifying known howls (trained dataset) and 75% accuracy in identifying unknown howls (test dataset). For the first time, the unknown wolf howls were classified successfully. Although the achievement is very significant in wolf vocalisation research, further accuracy is required for using them in the population estimation model. Training the model with more howls and verifying them with a different set of test data might increase its reliability. For these, a continuous recording of captive individuals and recordings from free-ranging collared wolves for an extended period is essential. The howling behaviour of Indian wolves has never been studied. Therefore, understanding the howling behaviour of the Indian wolf was the key to designing a howl survey methodology for population estimation. I studied the howling behaviour of collared and non-collared free-ranging wolves through the response pattern of the active howl survey. I found a disparity in their howl response - based on the distance to villages. In the low disturbed East-Maharashtra (EM), wolves mostly avoid responding to howling surveys (HS) if done within 1200 meters of villages [Response Rate(RR)=0.03±0.021], but they do respond once it is done far from villages (>1200m)[RR=0.226±0.075]. In high human dense West-Maharashtra (WM), wolves showed high RR within 1200 meters from the villages (RR=0.148±0.031). But the RR within 500 meters from villages is less as howling near villages might owe to easy detection. The collared wolf data showed significantly high RR (0.635±0.067) in their home-range core but low RR if the core area is close to a village. Therefore howling too close to the village is disadvantageous, although their tolerance for responding to HS has increased in the human-dominated landscape. The extent of the village may increase further with development, which will leave fewer areas for the wolf to defend territory with a long-range howl. The wolves might behaviourally adapt to a human-modified landscape by reducing their howling intensity. Adaptation in a fragmented habitat may save the wolves from extinction, but the repercussions of the fundamental behavioural alteration might adversely impact wolf behaviour and the ecological cascade. Whereas ecologists are mainly concerned with the extinction of species, the study highlights the vulnerability of fundamental behaviour of a keystone species attributed to human-induced contemporary evolution. Based on the vocalisation behaviour, I found that a howl survey should be done during their pre-denning season (November-December). Additionally, wind speed is low during this period. The best grid size for a systematic grid howl sampling is 1.7 × 1.7 km2. A 30watt speaker should be used for an active howl survey with 3-5 trials. This study provides the crucial guideline for a howling survey in Indian conditions. Based on these criteria, a howl survey was designed for four districts of Maharashtra. Maximum Entropy Probably Distribution (Maxent) was used for delineating the potential wolf habitats, and 12250 km2 effective wolf habitat was found. A newly triple observer-based howl survey method was introduced, I obtained a relatively high howl response (seven out of twenty-five howl surveys) in randomly selected grids. I used ‘redetection’ in different points in space instead of using individual ‘recapture’ with time. Through my pilot study, I found Indian wolf density is 3.65 individuals/100 km2 with a lower limit of 1.67 to an upper limit of 5.63 (95% CI). Although I do not have data on the population density of Indian wolves to compare, the data and its error range are comparable with the population density of Iberian wolves, i.e., 2.55 wolves/100 km2 (95% CI = 1.87–3.51) estimated by DNA (scat) sampling by López-Bao et al. (2018). The standard error might decrease further with an increase in sampling effort through the active howl survey. This methodology can be a guideline for using the active howling survey in the population estimation of wolves globally. Since wolf howls also possess individual information, incorporating this information in the future will help reduce the bias and heterogeneity in the population estimation model. Incorporating individual identification in the population estimation model will help generate additional details such as animal survival and home range. Regular population monitoring will help conserve and save this cryptic species before its population falls below a recovery level. Therefore, the study is a stepping stone towards using bioacoustics to estimate animal density and play a significant role in global wolf conservation.
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... during summer . The reason for a lower mobility during summer is 150 denning, when wolf activity concentrates on the den site and the breeding female does not 151 move far from the den Tsunoda et al., 2008). The intensity of 152 territorial marking predicts wolf reproduction (Llaneza et al., 2014). ...
How do seasonal changes in wolf defecation patterns affect scat detection probabilities of trained dog surveys
Preprint
Full-text available
  • Jun 2022
We used a detection dog and camera trapping (CT) to compare wolf scat detectability during winter and the reproductive season.
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... The variation in selection coefficients for anthropogenic and natural linear features was markedly larger in the rendezvous season compared to denning or winter. This pattern may be related to division of labour within the pack: wolves have different roles within the pack during the rendezvous season, with mothers generally staying closer to pups, and other pack members becoming more active to compensate [100]. Wolf packs tend to be more cohesive during winter with the pack functioning more as a single unit [85,101]. ...
Compensatory selection for roads over natural linear features by wolves in northern Ontario: Implications for caribou conservation
Article
Full-text available
Woodland caribou (Rangifer tarandus caribou) in Ontario are a threatened species that have experienced a substantial retraction of their historic range. Part of their decline has been attributed to increasing densities of anthropogenic linear features such as trails, roads, railways, and hydro lines. These features have been shown to increase the search efficiency and kill rate of wolves. However, it is unclear whether selection for anthropogenic linear features is additive or compensatory to selection for natural (water) linear features which may also be used for travel. We studied the selection of water and anthropogenic linear features by 52 resident wolves (Canis lupus x lycaon) over four years across three study areas in northern Ontario that varied in degrees of forestry activity and human disturbance. We used Euclidean distance-based resource selection functions (mixed-effects logistic regression) at the seasonal range scale with random coefficients for distance to water linear features, primary/secondary roads/railways, and hydro lines, and tertiary roads to estimate the strength of selection for each linear feature and for several habitat types, while accounting for availability of each feature. Next, we investigated the trade-off between selection for anthropogenic and water linear features. Wolves selected both anthropogenic and water linear features; selection for anthropogenic features was stronger than for water during the rendezvous season. Selection for anthropogenic linear features increased with increasing density of these features on the landscape, while selection for natural linear features declined, indicating compensatory selection of anthropogenic linear features. These results have implications for woodland caribou conservation. Prey encounter rates between wolves and caribou seem to be strongly influenced by increasing linear feature densities. This behavioral mechanism–a compensatory functional response to anthropogenic linear feature density resulting in decreased use of natural travel corridors–has negative consequences for the viability of woodland caribou.
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... Alternatively, wolves may use linear features to facilitate movement during territory monitoring, scent marking, travelling to and from rendezvous sites, among habitat patches, or den sites, without influencing hunting behaviours (Mech & Boitani 2003;Tsunoda et al. 2009;Giuggioli, Potts & Harris 2011). While movement rates increased when wolves used linear features, the time spent on these features was low, potentially limiting their effect on behaviours such as hunting. ...
Faster and farther: Wolf movement on linear features and implications for hunting behaviour
Article
Full-text available
1. Predation by grey wolves Canis lupus has been identified as an important cause of boreal woodland caribou Rangifer tarandus caribou mortality, and it has been hypothesized that wolf use of human-created linear features such as seismic lines, pipelines and roads increases movement, resulting in higher kill rates. 2. We tested whether wolves select linear features and whether movement rates increased while travelling on linear features in north-eastern Alberta and north-western Saskatchewan using 5-min GPS (Global Positioning System) locations from twenty-two wolves in six packs. 3. Wolves selected all but two linear feature classes, with the magnitude of selection depend- ing on feature class and season. Wolves travelled two to three times faster on linear features compared to the natural forest. Increased average daily travelling speed while on linear fea- tures and increased proportion of steps spent travelling on linear features increased net daily movement rates, suggesting that wolf use of linear features can increase their search rate. 4. Synthesis and applications. Our findings that wolves move faster and farther on human- created linear features can inform mitigation strategies intended to decrease predation on woodland caribou, a threatened species. Of the features that can realistically be restored, miti- gation strategies, such as silviculture and linear deactivation (i.e. tree-felling and fencing), should prioritize conventional seismic lines (i.e. cleared lines used for traditional oil and gas exploration) and pipelines, as they were selected by wolves and increased travelling speed, before low-impact seismic lines.
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... Preference for forest roads decreases with increasing road density within wolf home ranges (functional response, Houle et al. 2010). Reproducing wolves and their pups are likely most vulnerable to humans and other large predators during the summer season because movements are centered round a den site and later rendezvous sites (Jędrzejewski et al. 2001;Schmidt et al. 2008;Tsunoda et al. 2009). Survival of both adults and pups may be severely affected if these sites are detected. ...
Behavioral responses of wolves to roads: Scale-dependent ambivalence
Article
Full-text available
  • Oct 2014
Full text access here: http://beheco.oxfordjournals.org/content/early/2014/08/20/beheco.aru134.full Throughout their recent recovery in several industrialized countries, large carnivores have had to cope with a changed landscape dominated by human infrastructure. Population growth depends on the ability of individuals to adapt to these changes by making use of new habitat features and at the same time to avoid increased risks of mortality associated with human infrastructure. We analyzed the summer movements of 19 GPS-collared resident wolves (Canis lupus L.) from 14 territories in Scandinavia in relation to roads. We used resource and step selection functions, including >12000 field-checked GPS-positions and 315 kill sites. Wolves displayed ambivalent responses to roads depending on the spatial scale, road type, time of day, behavioral state, and reproductive status. At the site scale (approximately 0.1 km2), they selected for roads when traveling, nearly doubling their travel speed. Breeding wolves moved the fastest. At the patch scale (10 km2), house density rather than road density was a significant negative predictor of wolf patch selection. At the home range scale (approximately 1000 km2), breeding wolves increased gravel road use with increasing road availability, although at a lower rate than expected. Wolves have adapted to use roads for ease of travel, but at the same time developed a cryptic behavior to avoid human encounters. This behavioral plasticity may have been important in allowing the successful recovery of wolf populations in industrialized countries. However, we emphasize the role of roads as a potential cause of increased human-caused mortality.
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Denning phenology and reproductive success of wolves in response to climate signals
Article
Full-text available
Arctic and boreal ecosystems are experiencing rapid changes in temperature and precipitation regimes. Subsequent shifts in seasonality can lead to a mismatch between the timing of resource availability and species' life-history events, known as phenological or trophic mismatch. Although mismatch has been shown to negatively affect some northern animal populations, longer-term impacts across large regions remain unknown. In addition, animals may rely on climate cues during preceding seasons to time key life history events such as reproduction, but the reliability of these cues as indicators of subsequent resource availability has not been examined. We used remote sensing and gridded spatial data to evaluate the effect of climate factors on the reproductive phenology and success of a wide-ranging carnivore, the gray wolf (Canis lupus). We used GPS location data from 388 wolves to estimate den initiation dates (n = 227 dens within 106 packs) and reproductive success in eight populations across northwestern North America from 2000-2017. Spring onset shifted 14.2 days earlier, on average, during the 18-year period, but the regional mean date of denning did not change. Preceding winter temperature was the strongest climatic predictor of denning phenology, with higher temperatures advancing the timing of denning. Winter temperature was also one the strongest and most reliable indicators of the timing of spring onset. Reproductive success was not affected by timing of denning or synchrony with spring onset, but improved during cooler summers and following relatively dry autumns. Our findings highlight a disconnect between climate factors that affect phenology and those that affect demography, suggesting that carnivores may be resilient to shifts in seasonality and yet sensitive to weather conditions affecting their prey at both local and regional scales. These insights regarding the relationship between climate and carnivore demography should improve predictions of climate warming effects on the highest trophic levels.
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Program ochrony wilka Canis lupus w Polsce
Research
Full-text available
  • May 2017
Krajowa strategia ochrony wilka warunkująca trwałość gatunku w Polsce
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Accuracy of Radiotelemetry to Estimate Wolf Activity and Locations
Article
Full-text available
We analyzed radiotracking data from wolves (Canis lupus) in the Bia l / owie z . a Forest, Poland, to determine (1) differences between methods of estimating mean wolf activity and daily activity patterns, (2) whether activity estimated by changes in signal strength is dependent on the distance between the radiotracked wolf and the track -er, (3) radiotracker influence on wolf activity estimates, and (4) accuracy of radio locations. Daily patterns of wolf activity estimated by changes in signal strength, movements, and activity sensors were similar. However, the mean time spent active estimated by changes in signal strength (55%of the time) was higher than those estimated by movements (34%) or sensors (32%). We obtained the most accurate estimates of activity by a combination of move -ment, signal strength, and sensor data (43%) or by combining movement and signal strength data (44%). Activity estimated by changes in signal strength did not vary with the distance between radiotracked animals and radio -trackers. The trackers had no detectable influence on activity and movements of wolves when the tracker-to-wolf distance was >200 m. There was a small but not significant influence if trackers were <200 m away during the day. The mean radiotracking error was 194 m (95%CI: 157 – 231 m). We recommend that data on movements always be included in estimates of mean time spent active because activity sensors lead to underestimates and changes in sig -nal strength to overestimates. Distances traveled obtained by radiotracking should not be regarded as minimal dis -tances traveled, since the likelihood that they are overestimated or underestimated depends on the relation between the accuracy of radio locations and the mean distance that an animal travels per radiotracking interval. JOURNAL OF WILDLIFE MANAGEMENT 66(3):859–864
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Human impact on wolf activity in the Bieszczady Mountains, SE Poland
Article
Full-text available
  • Jun 2007
until recently. The radio tracked wolves of three packs moved throughout the day with one major peak around dawn. Wolves avoided the area around main public roads more at night (up to a distance of 1.5 km) than in the day (up to 0.5 km). Wolves avoided a 0.5-km area around secondary public roads and paved forest roads both at night and in the day but did not avoid the surroundings of settlements. As compared with other studies, wolves in this study were the least nocturnal although human density was the highest. We conclude that human activity is unlikely to be the reason for nocturnal activity in wolves.
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Regurgitative food transfer among wild wolves
Article
Full-text available
  • Nov 1999
Few studies of monogamous canids have addressed regurgitation in the context of extended parental care and alloparental care within family groups. We studied food transfer by regurgitation in a pack of wolves on Ellesmere Island, North West Territories, Canada, during 6 summers from 1988 through 1996. All adult wolves, including yearlings and a post-reproductive female, regurgitated food. Although individuals regurgitated up to five times per bout, the overall ratio of regurgitations per bout was 1.5. Pups were more likely to receive regurgitations (81%) than the breeding female (14%) or auxiliaries (6%). The breeding male regurgitated mostly to the breeding female and pups, and the breeding female regurgitated primarily to pups. The relative effort of the breeding female was correlated with litter size (Kendall's tau = 0.93, P = 0.01).
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Daily movements and territory use by radio-collared wolves ( Canis lupus ) in Bialowieza Primeval Forest in Poland
Article
Full-text available
  • Nov 2001
Wolves (Canis lupus) (9 females and 2 males from 4 packs), were radio-tracked in a large Polish woodland in Bialowieża Primeval Forest in 1996-1999. Based on 360 days of radio tracking with locations taken at 30- or 15-min intervals, daily movement distances (DMDs) of wolves and their utilization of territories were analyzed. DMDs averaged 22.1 km for females and 27.6 km for males. In reproductive and subadult females, DMDs varied seasonally, with the shortest daily routes in May and the longest in autumn-winter. Little seasonal variation was observed in nonbreeding and unsuccessfully breeding adult females. An adult male covered the longest DMDs in February (mating season). The mean speed of travelling wolves was 2.2 km/h. Wolves' hunting activity affected the length and speed of their movements, both of which were higher before than after a kill was made. With growing abundance of prey, DMDs of wolves became shorter. Snow cover and rainfall had a negligible effect on wolf travel. The mean straight-line distance between consecutive daily locations (SLD) was 4.4 km, i.e., on average, 21% of the actual route covered by wolves. Daily ranges utilized by wolves averaged 21.4 km2, or 9% of the whole territory. Variation in SLDs and daily ranges was shaped predominantly by mating, breeding, and pup rearing. The pattern of territory use by wolves differed between seasons. In spring-summer, their movements concentrated around the breeding den and rendezvous sites, and the areas used on consecutive days overlapped extensively. In autumn-winter, wolves moved widely and utilized their territory in a rotational way, returning to the same parts every 6 days, on average. Rotational use is related to intense patrolling and defense of territory, but may also help wolves to avoid behavioral depression of prey availability.
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Daily and seasonal variation in wolf activity in the Bieszczady Mountains, SE Poland
Article
Full-text available
  • Mar 2009
Wolf activity varies considerably among differentstudies that explained activity patterns by factors suchas human activity, breeding status or prey availability(Fancy and Ballard 1995; Vila` et al. 1995; Ciucci et al.1997; Theuerkauf et al. 2003; Kusak et al. 2005; Chavezand Gese 2006; Theuerkauf et al. 2007). However, it islikely that the high variability of wolf activity patterns isa result of their ability to react to various environmentalconditions (Packard 2003). Activity of individual wolvesmay also vary among days, switching from being diurnalto nocturnal. The purpose of our study was to examinethe variability of wolf activity between days and seasonsand to determine the most important factors thatinfluence activity in the Bieszczady Mountains, Poland.The study was conducted in the Bieszczady Moun-tains (Polish Carpathians) over an area of about1000km
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Movement Patterns of Barren-Ground Wolves in the Central Canadian Arctic
Article
Full-text available
  • Aug 2001
We collected information on the movement patterns of wolves (Canis lupus) captured within a 30,000-km2 area in the Northwest Territories and western Nunavut. Currently, diamond mining and road construction are occurring in the area used by these migratory wolves for denning. During summers of 1997 and 1998, 23 wolves in 19 different packs were captured and fitted with collar-mounted satellite transmitters. Areas used by these wolves varied seasonally and seemed to correspond to movements of migratory caribou (Rangifer tarandus). Annual home-range sizes (95% minimum convex polygon), averaged 63,058 km2 ± 12,836 SE for males and 44,936 ± 7,564 km2 for females. Wolves began to restrict movements around a den site on the tundra by late April. They did not depart from their summer ranges until late October, after which they followed caribou to their wintering grounds. Straight-line distances from the most distant location on the winter range to the den site averaged 508 ± 26 km during 1997-1998 and 265 ± 15 km in 1998-1999 (P < 0.01). Home range in summer averaged 2,022 ± 659 km2 for males and 1,130 ± 251 km2 for females. No difference was detected between sexes or years. All but 2 of 15 wolves returned to <25 km of a previous den, and 2 wolves returned to the same den site. We believe that human activities that disturb or displace denning wolves, or that alter the distribution or timing of caribou movements, will have negative affects on reproductive success of wolves.
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Denning behaviour of non-gravid Wolves, Canis lupus
Article
  • Jan 1996
Wild Wolves (Canis lupus) that had produced pups in earlier years but were not currently pregnant, and ovariectomized captive Wolves, dug dens during and after the whelping season even though they produced no pups. These observations suggest that den digging is not a function of pregnancy or of ovarian estrogen or progesterone. We hypothesize that increasing prolactin in spring elicits or mediates den-digging behavior.
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Differential use of a Wolf, Canis lupus, pack territory edge and core
Article
Based on 418 radio-locations of a Minnesota Wolf pack, Wolves were found at significantly fewer locations per area in the outer 2 km of the territory than in the core. This finding supports an hypothesis that buffer zones exist between pack territories and may explain why prey survive longer there.
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