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Seasonal and sexual variation in diel activity rhythms of pine marten Martes martes in the Bialowieza National Park (Poland)

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From 1991–1996, the activity rhythms of 14 radio-collared pine martens Martes martes (Linnaeus, 1758) (6 males and 8 females) were studied in the pristine deciduous and mixed forests of the Białowieża National Park. Tracking data (5823 h) indicated that the activity rhythms of pine martens varied between sexes and seasons. In spring, male activity peaked at 20.00–00.00 h, whereas in summer and autumn-winter, activity was bimodal, peaking at 18.00–22.00 h and 02.00–04.00 h. Female activity in spring was more evenly distributed than that of males, but in summer their activity peaked at 20.00–00.00 h, while in autumn-winter females had a bimodal rhythm with peaks at 18.00–20.00 h and 02.00–06.00 h. In breeding females, activity rhythms changed in the course of pregnancy and nursing. On average, martens started their activity 73±209 (SD) min before sunset and finished 87±245 min after sunrise. Females became active earlier than males but both sexes terminated activity at the same time. For both males and females the daily activity rhythm was not related to the diurnal course of temperature. Key words Martes martes-activity rhythms-thermal stress-Białowieża National Park
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Seasonal and sexual variation
in diel activity rhythms of pine marten Martes martes
in the Bia³owie¿a National Park (Poland)
Zalewski A. 2001. Seasonal and sexual variation in diel activity rhythms of pine marten
Martes martes in the Bia³owie¿a National Park (Poland). Acta Theriologica 46: 295–304.
From 1991–1996, the activity rhythms of 14 radio-collared pine martens Martes
martes (Linnaeus, 1758) (6 males and 8 females) were studied in the pristine deciduous
and mixed forests of the Bia³owie¿a National Park. Tracking data (5823 h) indicated
that the activity rhythms of pine martens varied between sexes and seasons. In spring,
male activity peaked at 20.00–00.00 h, whereas in summer and autumn–winter, activity
was bimodal, peaking at 18.00–22.00 h and 02.00–04.00 h. Female activity in spring was
more evenly distributed than that of males, but in summer their activity peaked at
20.00–00.00 h, while in autumn–winter females had a bimodal rhythm with peaks at
18.00–20.00 h and 02.00–06.00 h. In breeding females, activity rhythms changed in the
course of pregnancy and nursing. On average, martens started their activity 73±209
(SD) min before sunset and finished 87 ± 245 min after sunrise. Females became active
earlier than males but both sexes terminated activity at the same time. For both males
and females the daily activity rhythm was not related to the diurnal course of
Mammal Research Institute, Polish Academy of Sciences, 17-230 Bia³owie¿a, Poland,
Key words:Martes martes, activity rhythms, thermal stress, Bia³owie¿a National Park
Animals adapt their circadian activity to diurnal variation in their environment,
food availability, predation risk, and reproduction constraints (Aschoff 1964, Daan
and Aschoff 1982). Each of these factors plays a different role relative to season but
two factors have often been used as an explanation of predator’s activity rhythms:
predator-prey synchronisation and predation risk. It is assumed that increased
predator-prey synchronisation of activity leads to increased predator hunting
success. Many studies focus on this factor as an explanation of predators activity
rhythms (Ables 1969, Gerell 1969, Schuh et al. 1971, Curio 1976, Koop and
Velimirov 1982, Zielinski et al. 1983, Ferguson et al. 1988, Lodé 1995, Lariviere and
Messier 1997). The pine marten Martes martes (Linnaeus, 1758) prey mainly on
small rodents but the composition of their diet changes between seasons (Grakov
1981, Jêdrzejewski et al. 1993). In Bia³owie¿a National Park, bank voles Clethrio-
Acta Theriologica 46 (3): 295–304, 2001.
PL ISSN 0001–7051
nomys glareolus are the preferred prey species throughout the year and constitute,
on average, 32% of the biomass consumed by martens (Jêdrzejewski et al. 1993).
Avoidance of predation by small and medium-sized predators themselves has also
been suggested as the only adequate explanation of patterns of activity (Geffen and
Macdonald 1993, Drew and Bissonette 1997). The pine marten is a medium-sized
mustelid and could be killed by many predators and raptors (Korpimäki and
Norrdahl 1989, Lindström et al. 1995, Okarma et al. 1997).
Seasonal variation of activity patterns, however, was strongly affected by
variation of weather and reproduction constrains. Since the pine marten has a thin,
elongated body conductive to rapid heat loss (Iversen 1972), its activity rhythm
may be influenced by variation in diel temperature. At low temperature martens
curtailed their activity, however, they were mostly active at night (Zalewski 2000).
In winter, the American marten Martes americana was active during daylight
hours when temperature was higher than during the night (Thompson and Colgan
1991). Male pine martens are 33% larger than females. As a consequence females
need to reduce their activity during periods of cooler temperature much more than
males. In winter, the smaller females need to be active more often due to higher
possibility of starvation but they are active for a shorter duration (Zalewski 2000).
It could be expected that females would be active more evenly in the day.
The different investment in parental care by males and females can also affect
activity, and sexual differences in activity pattern are likely to occur, especially in
spring and summer. At this time females invest their energy in parental care in
order to maximise breeding success. In spring, females leave their cubs for short
periods, but duration of activity did not differ from that in males (Zalewski 2000).
Thus female activity is more evenly distributed during the day, which make them
more active in daylight hours. This could result in females running a higher
predation risk by raptors.
In this paper, I examine circadian rhythms of pine marten activity over a five-
-year period in the last remnant of pristine deciduous and mixed forests in the
European lowlands, where the pine martens exist amongst very rich communities
of prey and other predators (Jêdrzejewska and Jêdrzejewski 1998). The area has
not been exploited for timber and the level of human penetration is very low.
Therefore, human impact on marten behaviour is negligible. In this paper, I
analysed: (1) seasonal variation in activity rhythms of pine marten; (2) differences
in the daily activity rhythms between male and female; (3) variation of females
activity rhythms in pregnancy and nursing.
Study area
The study was conducted in an area of north-east Poland (52°43’N, 23°54’E) within the strict
reserve of Bia³owie¿a National Park (47.5 km2– BNP). The Park is part of a large primeval woodland
covering over 1250 km2, in which old-growth forest areas are dominated by oak-lime-hornbeam stands
(44.4% of area) comprising hornbeam Carpinus betulus, oak Quercus robur and lime Tilia cordata,as
well as scattered spruce Picea abies. Two other main forest types are mixed coniferous (dominated by
296 A. Zalewski
spruce and pine Pinus silvestris) and ash-alder (dominated by black alder Alnus glutinosa and ash
Fraxinus excelsior). More detailed information on the vegetation of BNP is given by Faliñski (1986).
Old-growth forests of BNP are characterised by various age of trees (mean age of tree stands is 130
years), the presence of snags and downed logs of large diameter, and small gaps in the canopy. Human
penetration (mainly pedestrian tourists) of the National Park is very low and only occurs in the
south-western part of the study area.
The climate is transitional between continental and Atlantic types but continental features prevail
(Olszewski 1986). During the study (1991–1996), January or February were the coldest months, with
average monthly temperature reaching as low as –8.5°C, and a maximum snow cover of 63 cm. The
warmest month was July, with average monthly temperature reaching a high of 22.5°C. More detailed
information about the area as well as maps of the terrain are given by Jêdrzejewska and Jêdrzejewski
Material and methods
From 1991 to 1996, 6 male and 8 female pine martens were captured in box traps and equipped
with radio-collars (AVM or Lotek; 12–25 g). Martens were located at least once daily and on some days
they were monitored continuously for a time span of 4–24 h. During such continuous radiotracking,
locations were taken at 15-min intervals. Marten behaviour was categorised as active (frequently
switching pulse amplitude) or inactive (pulse amplitude unchanging). Since slight movements of the
animal in a resting site could have been confused with true locomotor activity, the activity was
recorded as occurring in the resting site if the marten’s spatial position did not change despite a slight
variation in the signal. In total, 23 294 activity fixes (5823 h) were recorded (15203 for males and 8091
for females). For analysis of the diel activity rhythms in each season, the total number of fixes during
2-hour periods were taken as 100 so active fixes were thus calculated as percentages. Pooled data from
three seasons: spring (16 March – 15 June), summer (16 June – 15 October) and autumn–winter (16
October – 15 March) were analysed. The first date upon which natal dens were located was accepted as
parturition date, although such estimates may be 2–4 days after the actual birth.
Circadian rhythms of marten activity were also correlated with diel course of ambient temperature
measured in 1960, in oak-lime hornbeam forest of BNP at 0.2 m above ground (Olszewski 1986). In
1960, average temperature in January (–4.5°C) was within the range of mean January temperatures in
my study period (–0.9 to –8.5°C). The mean daily snow cover in 1960 was 6.2 cm, again within the
range recorded in 1991–1996 (1.7 to 20.7 cm).
Diel distribution of martens activity was not homogenous. Generally, activity
levels were higher between 18.00–06.00 h followed by a decline during daylight
hours (Fig. 1). In both sexes activity varied significantly between seasons (G-test
for homogeneity of percentages: males – Gfrom 25.5 to 38.0, p< 0.01, df = 11;
females – Gfrom 49.3 to 69.5, p< 0.001, df = 11). In spring, male activity peaked at
20.00–00.00 h (91 and 87% of active fixes) and was at its lowest at 08.00–14.00 h
(9–10%, Fig. 1). In summer, males displayed a bimodal activity rhythm with peaks
at 18.00–22.00 h (75 and 77% of active fixes) and 02.00–04.00 h (79% of active fixes;
Fig. 1). Daylight activity (minimum 10% between 12.00–14.00 h) was noticeably
higher in summer than during any other season. In autumn–winter, male martens
also displayed a bimodal rhythm of activity with a maximum of 50–60% of active
Activity rhythms of pine marten 297
298 A. Zalewski
Fig. 1. Circadian activity rhythms of male (n= 6) and female (n= 8) pine martens Martes martes
during spring (15 March–15 June), summer (16 June–15 October) and autumn–winter (16 October–15
March) in the Bia³owie¿a National Park (Poland) in 1991–1996. For each 2-hour period fixes when
martens were found active were calculated as percentage of all fixes taken at 15-min intervals. Total n
fixes in 2-h periods varied from 83 to 1741. Dark horizontal bars indicate the shortest and the longest
time from sunset to sunrise in each season.
Active out of a resting site (moving) Sunset to sunrise tim
Active in a resting site
12 16 20 004 08 12
12 16 20 00408 12
12 16 20 004 08 12 12 16 20 004 08 12
12 16 20 004 08 12
12 16 20 00408 12
Autumn - winter
Hours Hours
Percentage o
Males Females
In spring, female activity rhythms were more evenly distributed than those of
males. Their level of activity shown for 2-hour periods varied from 29% to 63%,
except between 04.00–10.00 h, when it was lower (13–18%, Fig. 1). In summer,
female activity peaked between 20.00–00.00 h (89–91% of active fixes). In autumn–
–winter, females, like males, had a bimodal activity rhythm, exhibiting a bigeminus
pattern (Aschoff 1966) with the first peak markedly higher than the second.
These data show different rhythms of activity in males and females, especially in
spring (spring: G= 62.4, p< 0.001; summer: G= 22.3, p< 0.05; autumn–winter:
G= 42.4, p< 0.05, df = 11 in each case). In spring, females were more active than
males between 10.00–16.00 h (Gfrom 4.5 to 17.9, p< 0.05, df = 1) but males were
significantly more active between 20.00–00.00 h (G= 8.2 and 12.5, p< 0.01,
df = 1; Fig 1). In summer, the activity rhythms of the sexes were closer than at any
other season but there were still significant differences between 22.00–00.00 h
(G= 6.9, p< 0.01, df = 1) and 02.00–04.00 h (G= 4.0, p< 0.05, df = 1). During
autumn– –winter, females decreased their activity to a much greater extent than
males between 20.00–00.00 h (G= 6.6 and 18.6, p< 0.05, df = 1) and 02.00–04.00 h
(G= 6.3, p< 0.05, df = 1) but were more active between 10.00–14.00 (G= 6.3 and
5.2, p< 0.05, df = 1; Fig. 1). Marten activity at resting sites was always of short
duration (on average 0.3–1.1% of all fixes in 2-h periods, ie 5–24 min/day) with the
exception of females in spring (on average 3.6% of fixes, ie 51 min/day; Fig. 1).
In all seasons daily ambient temperatures are lowest between 03.00–06.00 h.
The daily activity pattern of both males and females was not positively related to
the diurnal course of temperature (Kendall’s coefficient of rank correlation: for
males tfrom –0.27 to –0.30; for females tfrom 0.09 to 0.24; p> 0.05). However,
between 04.00 and 08.00 h, when ambient temperature was at its lowest, martens
often decreased their activity (especially females in spring).
For breeding females activity rhythms changed during the course of pregnancy
and nursing. Indeed, in the last month of her pregnancy, female 6 was even active
between 14.00–06.00 h (Fig. 2). During the first month of the cubs’ life, activity
rhythms of all lactating females showed two peaks and similarly all females
decreased their activity between 02.00 to 10.00 h. In the same period, however,
females were generally more active at their resting sites, especially between
06.00–22.00 h. Periods of activity at the resting site decreased during the second
month of the cub’s life and two females were again active outside the resting site
between 14.00–02.00 h (Fig. 2).
On average, martens started their activity 73 min (SD ± 209) before sunset and
terminated it 87 ± 245 min after sunrise. However, the onset and cessation of
activity was highly variable according to both sex and season (Table 1). Overall
females started their activity earlier than males (two-way Kruskal-Wallis ANOVA:
H= 6.4 , p< 0.01) and both sexes started activity earlier in summer and later in
winter (H= 45.9, p< 0.001). Males and females both terminated activity at the
same time (H= 0.05, ns) with termination time later in summer than in winter
(H= 40.1, p< 0.001; Table 1).
Activity rhythms of pine marten 299
300 A. Zalewski
10 14 18 22 02 06 10
10 14 18 22 02 06 10 10 14 18 22 02 06 10
10 14 18 22 02 0610 10 14 18 22 02 0 610
10 14 18 22 02 06 10
10 14 18 22 02 06 10
6(15 Mar - 16 Apr 1993)
6(17 Apr - 18 May 1993)
(16 May - 17 Jun 1992)
(4 Apr - 5 May 1992)
(6 May - 7 Jun 1992)
(7 May - 8 Jun 1992)
(19 May - 20 Jun 1993)
Hours H
Active out of den
Active in den
First month after parturition
Second month after parturition
Percentage of fixes
Fig. 2. Circadian activity rhythms of female pine martens during their pregnancy, the first and the
second month after parturition in the Bia³owie¿a National Park. Total nfixes in 4-h periods varied
from 40 to 150.
The results show that the activity rhythms of pine martens varied significantly
between sexes and seasons. The differences between sexes could be due to
differences in the reproduction investment of both sexes and this was most evident
during spring. In this season, females care for their cubs (Grakov 1981). They leave
the cubs for short bouts because unattended cubs may be susceptible to thermal
stress (Frost and Krohn 1997). In spring, the activity of females was more evenly
distributed throughout the day than that of males. Similar behaviour has been
observed in females of other predator species (Paragi et al. 1994, Lariviere and
Messier 1997). The even distribution of female activity during this season was
apparently related to their care for young, thus producing more frequent activity
bouts in a day, although each bout was of shorter duration than those of males
(Zalewski 2000). However, both sexes were active for a similar duration (Zalewski
2000). Therefore, female activity started earlier in the day compared to males
because they often had breaks in their activity to care for their cubs. In spring,
females decreased their activity during the coldest part of the day (04.00–08.00 h)
returning to their den at that time. This was particularly marked during the first
month of a cub’s life. In spring, females were also active longer in dens than at any
other season, especially during the first month of cub’s life. Females choose
breeding sites in high cavities of trees (Zalewski 1997), where the risk of predation
is low. These sites are accessible only to other martens. It is possible that reversal of
activity between males and females is affected by male predation on cubs. However,
there is no evidence of intraspecific predation. In America, male martens were
Activity rhythms of pine marten 301
Table 1. Daily onset and cessation of activity period in relation to sunset and sunrise for pine martens
Martes martes in the Bia³owie¿a National Park, 1991–1996. Positive and negative values indicate time
(min) following and preceding sunset or sunrise, respectively. n– number of observation.
Sex of
Onset of activity in relation to sunset
Cessation of activity in relation to sunrise
martens nAverage (SD) nAverage (SD)
Males 17 +90 (178) 17 +113 (185)
Females 21 +210 (151) 16 +113 (275)
Males 27 +125 (137) 23 +224 (143)
Females 7 +250 (166) 14 +254 (170)
Males 16 –190 (208) 19 –76 (128)
Females 12 –55 (94) 11 –209 (292)
observed in breeding dens, scent marking or robbing the female of her prey, but not
killing cubs (Jones et al. 1997). In summer, both males and females synchronized
their activity pattern due to mating season.
In the Bia³owie¿a National Park, the daily activity pattern of both males and
females were not related to the diurnal course of temperature during any season. In
Ontario, American martens experiencing severe weather conditions remained in
their resting sites all night and confined much of their activity to daylight hours
when temperature were highest (Thompson and Colgan 1991). In California, the
winter activity of martens was distributed more homogeneously throughout 24 h
cycles (Zielinski et al. 1983, Martin 1987). In BNP, pine martens were only active
during the night in the winter, despite the fact that temperatures were lowest then.
In winter, however, females decreased their activity in colder periods of the night.
Other studies have recorded only nocturnal activity amongst pine marten popu-
lations (Marchesi 1989). Drew and Bissonette (1997) suggest that the avoidance of
daylight by American martens was due to higher predation risk. This explanation
cannot be applied to pine martens in BNP. European pine martens are killed by the
lynx Lynx lynx, red fox Vulpes vulpes or large raptors (Nyholm 1970, Korpimäki
and Norrdahl 1989, Pulliainen 1981, Lindström et al. 1995, Okarma et al. 1997). In
BNP, most of raptors are absent in winter (Pugacewicz 1996), while lynx and red
fox are mainly nocturnal (Schmidt 1999, R. Kowalczyk, pers. comm). In winter,
therefore, predation risks for martens seems to be even higher at night.
It is often assumed that predators synchronise their activity rhythms with that
of their main prey (Ables 1969, Mikkola 1970, Curio 1976, Zielinski et al. 1983,
Weber et al. 1994, Lodé 1995). Yellow-necked mice Apodemus flavicollis were active
at night (Buchalczyk 1964, Wójcik and Wo³k 1985) and the activity pattern of pine
martens correlated with that of this species. However, martens consistently preyed
on mice less than could have been expected from the proportion of mice in the total
biomass of forest rodents determined by trapping (Jêdrzejewski et al. 1993). This
suggests that the activity rhythms of both predator and prey are not related but
depend on other factors and synchronisation of predator-prey patterns of activities
do not implicate predator specialisation in this prey. Bank vole are the preferred
prey species by pine marten (Jêdrzejewski et al. 1993). In spring and summer, bank
voles showed activity patterns with several peaks throughout the day and night
(Buchalczyk 1964, Wójcik and Wo³k 1985). During these two seasons, the activity
patterns of both male and female pine martens did not synchronise with those of
bank voles. In autumn, however, bank voles’ activity displayed two peaks: at
18.00–20.00 h and 04.00–08.00 h (Buchalczyk 1964, Wójcik and Wo³k 1985). This
coincided with the circadian rhythm of pine marten activities (especially males) at
this time of year.
In conclusion, reproduction clearly affected the intersexual variation in the
circadian rhythm of marten activity. However, marten activity behaviour does not
clearly relate to the activity rhythms of its main prey nor to thermal stress. There is
no evidence either to suggest that martens are active at night to avoid predation.
302 A. Zalewski
Acknowledgements: I wish to thank the technical staff of MRI (E. Bujko and K. Zub), students of
Farnborough College of Technology (England), Warsaw University, the Jagiellonian University in
Kraków, and Earthwatch Research Corps for their help in field work. Special thanks go to
Dr B. Jêdrzejewska for her help in the preparation of this manuscript. My thanks to Prof S. W.
Buskirk and Z. Pucek for their comments on an earlier draft. The study was supported partly by the
Mammal Research Institute PAS budget and partly by a KBN 6 P205 080 06 grant.
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Zielinski W. J., Spencer W. D. and Barrett R. H. 1983. Relationship between food habits and activity
patterns of pine martens. Journal of Mammalogy 64: 387–396.
Received 1 June 2001, accepted 22 June 2001.
304 A. Zalewski
... Specifically, in winter, low temperatures cause high thermal stress, and snowfall imposes travel costs and reduces hunting efficiency (Zhou et al. 2011;Willebrand et al. 2017), greatly affecting energy loss for carnivores (Bartoń and Zalewski 2007;Pozzanghera et al. 2016;Mustonen and Nieminen 2018). Previous studies have shown that carnivores, such as Canidae and Martes sp., limit their activity at low temperatures and/or heavy snowfall to reduce energy loss, and increase their activity at high temperatures and/or little snowfall in winter (e.g., Ables 1969;Zalewski 2001;Kauhala et al. 2007). These results indicate that in severe winters, multiple species can be more active simultaneously under similar weather conditions (i.e., high temperature and/ or little snowfall) to reduce the energy costs. ...
... It has also been confirmed that winter weather conditions affect the activity of red foxes, raccoon dogs, and Martes sp. (e.g., Ables 1969;Zalewski 2001;Kauhala et al. 2007). Therefore, these three species are suitable targets for testing our hypotheses. ...
... RF, RD, and JM represent the red fox, raccoon dog, and Japanese marten, respectively RF red fox, RD raccoon dog, JM Japanese marten + Temporal niche overlap; − temporal niche partitioning; Blanks represent there was neither overlap nor partitioning Drygala 2013). Zalewski (2001) have reported that Martes sp. were less active during cold winter nights. In addition, deep and fluffy snow limits the travel of red foxes and raccoon dogs (Ables 1969;Kauhala et al. 2007;Pozzanghera et al. 2016) and reduces the foraging efficiency of red foxes and Martes sp. ...
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Temporal niche partitioning may be influenced not only by interspecific competition, but also by weather conditions. Decreased food availability and dietary overlap between species can increase the degree of interspecific competition during winter, thereby promoting temporal niche partitioning. However, multiple species can be simultaneously active under similar weather conditions (high temperature and little snowfall) in winter to reduce energy costs and increase temporal niche overlap. In this study, we aimed to determine the degree of temporal niche partitioning among red foxes (Vulpes vulpes), raccoon dogs (Nyctereutes procyonoides), and Japanese martens (Martes melampus), and its variation with seasonal climate change in terms of interspecific competition and weather conditions. We obtained data on the target species through a camera-trap survey conducted in a heavy snowfall area in northeastern Japan. We analyzed the degree of temporal niche partitioning based on diel activity overlap, co-occurrence rates per night, and behavioral avoidance within 1 or 2 h. We also evaluated the relationship between the presence or absence of activity per night and nightly weather conditions (temperature, precipitation (snowfall), moonlight). We observed a high degree of temporal niche overlap among the three species. In particular, the degree of temporal niche overlap was higher in winter than that in other seasons because the activity of the three species was greatly affected by low temperatures and snowfall in winter. As a winter survival strategy, coping with the weather may be more important than avoiding competition. Our results conflict with the hypothesis predicting temporal niche partitioning in winter, suggesting that weather effects can be an important factor in varying temporal niche partitioning among carnivores. Significance statement This study revealed that the temporal niche overlap of three carnivores in northeastern Japan was higher in winter than that in other seasons because they were simultaneously active under similar weather conditions (high temperature and little snowfall) in winter to reduce energy costs. This indicates that coping with the weather may be more important than avoiding interspecific competition as a winter survival strategy. Our results conflict with the hypothesis predicting temporal niche partitioning among carnivores in winter due to restricted food resources and dietary overlap. This suggests that the different degrees of weather effects can be an important factor in varying temporal niche partitioning among carnivores. Because extreme changes in weather conditions such as extremely high temperatures, blizzards, and heavy rains, have occurred worldwide in recent years, weather conditions may significantly affect the niche partitioning among carnivores distributed in different environments worldwide.
... When these martens live in sympatry with several other carnivore species (e.g., the red fox Vulpes vulpes and the European badger Meles meles), stone martens are mostly nocturnal, whereas pine martens show a cathemeral or diurnal pattern (Fonda et al., 2017;Monterroso et al., 2014;Torretta et al., 2017). Zalewski (2001) showed that, in Central Europe (Białowieża Primeval Forest), the activity rhythms of the pine marten change seasonally, with daytime bouts increasing when cubs are present and with strictly nocturnal habits in winter. Where large competitors (e.g., the lynx Lynx lynx, the red fox, and large-sized raptors) are present, avoidance of daylight may result from predation risk perception by martens (Drew & Bissonette, 1997;Zalewski, 2001). ...
... Zalewski (2001) showed that, in Central Europe (Białowieża Primeval Forest), the activity rhythms of the pine marten change seasonally, with daytime bouts increasing when cubs are present and with strictly nocturnal habits in winter. Where large competitors (e.g., the lynx Lynx lynx, the red fox, and large-sized raptors) are present, avoidance of daylight may result from predation risk perception by martens (Drew & Bissonette, 1997;Zalewski, 2001). Because of their semi-retractable claws, pine martens widely use trees and canopies ranging throughout woodland areas, whereas tree cavities are required for reproduction and resting sites (Birks et al., 2005). ...
... As well, both Fonda et al. (2017) and Torretta et al. (2017) detected the same activity peaks in the early morning and around sunset, respectively, in the Carnic Prealps (North-Eastern Italy) and Liguria (North-Western Italy), both areas characterized by a rich carnivore guild. A slight increase in diurnal activity has been detected in spring, in line with all previous studies on the activity of pine martens (Clevenger, 1992;Fonda et al., 2017;Monterroso et al., 2014;Torretta et al., 2017;Zalewski, 2001). Bouts of diurnal activity may represent a female prerogative in early spring when nights start to be shorter concerning the winter months and when cubs are present (Zalewski, 2001). ...
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Behavioral interference and interspecific competition shape the spatiotemporal behavior of carnivores, where intra-guild predation has been recorded as a strategy to limit competition. Very often, disentangling the effect of intra-guild effects from other ecological processes is challenging, if not impossible. This work aimed to assess the spatiotemporal behavior of the pine marten Martes martes in an island ecosystem without any intra-guild predation. Using an intensive camera-trap survey on Elba Island, Italy, we estimated occupancy, detection probability, and locomotor rhythms of the species. The pine marten occurred throughout the study area, showing a tendency to avoid urban areas during the high tourist season. Occupancy was higher in colder months and lowered in autumn, following opposite fluctuations of recorded human presence. Conversely, the detection probability remained low throughout the year, confirming the pine marten elu-siveness. With other studies conducted in sympatry with other carnivores or predators , Elba Island martens showed locomotor ground activity mostly at dawn and dusk throughout the year, with an increasing diurnal locomotor activity in spring, when cubs are present. The spatiotemporal behavior of the pine marten in Elba island did show little variation across seasons. It was similar to those reported where potential predators were present, suggesting the intra-guild predation not playing a significant role in shaping the spatiotemporal activity of pine martens. K E Y W O R D S activity rhythms, island ecosystem, Martes martes, occupancy modeling, wildlife camera-trapping
... However, there is also a great deal of intraspecific variation in activity patterns, which may result from intersexual differences based on strong sexual dimorphism in body size (e.g. Marcelli et al., 2003;Begg et al., 2016) or reproductive constraints (Zalewski, 2001;Kolbe & Squires, 2007;Begg et al., 2016). In addition, important biogeographical variation between populations may be explained by climatic conditions and differential availability of food resources (Kowalczyk et al., 2003;Zalewski et al., 2004), while at the individual level meteorological factors (especially temperature) and food availability may influence the time and energy animals invest in their activities (Zielinski et al., 1983;Zalewski, 2000;Zalewski et al., 2004;Baghli & Verhagen, 2005). ...
In Europe, the stone marten, Martes foina , is one of the main carnivore species to inhabit urban areas. While these environments are generally resource rich, they also present a range of anthropogenic stresses, such as human persecution or road traffic, which have the potential to induce behavioural responses in urban wildlife. We radio‐tracked 12 stone martens in two towns in Luxembourg in order to determine how their activity (duration of the principal activity period, nightly activity duration) and movement (nightly movement distance, movement speed, nightly range) patterns were adapted to this environment. Stone martens displayed a more strictly nocturnal lifestyle than was known from studies on this species in more rural environments. We argue this to be a behavioural adaptation to reduce the rate of potential contact with humans. In fact, during long winter nights, emergence from dens took place long after sunset and return to dens intervened mostly before traffic picked up in the mornings. Furthermore, during long nights, marten peak activity was shifted to those parts of the night when human activity was at its lowest. On the other hand, stone martens were active during the entire dark period during short summer nights. Despite presumably higher resource availability and somewhat smaller territories compared to other studies, stone marten activity duration and movement distances were similar to those recorded in forest or rural populations elsewhere. Interestingly spring mobility was more pronounced in females than would be expected. We discuss these results in the context of territorial behaviour which, in urban areas, is likely to be driven by factors such as increased perceived intruder pressure.
... Para evitar posibles confusiones con la garduña (Álvares & Brito 2006, López-Martín 2007, Reig 2007, Barja 2017, cuya actividad es principalmente nocturna (Dudin & Georgiev 2015, Mangas 2017, Torretta et al. 2017, Roy et al. 2018, se muestreó en los meses en los que la marta presenta una mayor actividad diurna, entre finales de primavera y principios de otoño (Zalewski 2001, Petrov et al. 2016, Ruiz-González 2016, cuando los ejemplares están en celo y patrullan su territorio más intensamente (Zalewski 2000, Zalewski et al. 2004, Birks 2017. Para la asignación de las imágenes a una u otra especie de Martes, que pueden ser confundidas con facilidad incluso por observadores con experiencia, sobre todo en condiciones de luz escasa u observaciones fugaces (Blanco 1998, Barja 2017, se tuvieron en cuenta aspectos morfológicos, principalmente el color y la forma del babero, pero también el tamaño y posición relativa de las orejas, el aspecto general del individuo, o el color del pelaje y del rinario (Burki et al. 2010, Ruiz-González 2016, Birks 2017. ...
Pine marten Martes martes is a medium size mustelid that inhabits well-conserved forests of the northern Iberian Peninsula. Currently, the distribution and ecology of the species in the Iberian Peninsula is poorly known. In this work, 27 UTM 10x10km grids in the province of Ourense were sampled using camera-trapping techniques, with the aim of increasing knowledge about the distribution of the species in Galicia. The presence of pine marten was confirmed in 18 of the surveyed grids, which increases the species´ known distribution area by 21% in this community and by 62% in Ourense province, confirming camera-trapping specific survey as a highly effective method to know the distribution of elusive species such as the pine marten.
... таблицу). Наблюдаемый половой диморфизм соответствует индивидуальным предпочтениям куньих, заключающимся в разных энергетических потребностях и особенностях поведения полов [27][28][29]. Тот факт, что самки сильнее отдалялись от рек и ручьев намного дальше самцов, связан с возможной конкуренцией между ними. Во время неблагоприятных периодов года самки не заходят в буферные зоны или соседние участки ввиду возможной агрессии со стороны других особей и вынуждены удаляться от водоема на значительное расстояние. ...
... Therefore, at least during the summer, contact between the badgers and the vectors, which display crepuscular activity, is possible [21]. According to observed movement patterns, which involve various degrees of diurnal activity [22][23][24], martens and polecats would theoretically be more exposed to the vectors compared to badgers. Indeed, the relative frequency in beech martens seems to be higher compared to badgers. ...
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Background: Thelazia callipaeda (Spirurida, Thelaziidae) is a vector-borne zoonotic eye worm with a broad host spectrum. In Europe, it is an emerging threat, having greatly expanded its geographical distribution during the past two decades. In Romania, T. callipaeda has been previously reported in domestic and wild canids and felids. The aim of the present study was to assess the occurrence of T. callipaeda in mustelids in the country. Methods: Between March 2015 and April 2019, 77 road-killed mustelids (3 pine martens, Martes martes; 6 European polecats, Mustela putorius; 13 beech martens, Martes foina; and 55 European badgers, Meles meles) were examined by necropsy. If present, all ocular nematodes were collected and stored in absolute ethanol, for subsequent morphological and molecular identification. Results: Two animals were found to be infected with T. callipaeda: one European badger and one beech marten. The molecular analysis revealed a 100% nucleotide similarity to T. callipaeda haplotype h1 for all the sequenced specimens. Conclusions: To our knowledge, the present study demonstrates for the first time the occurrence of T. callipaeda in mustelids from Romania, records the easternmost locality of the parasite in Europe, and represents the first report of T. callipaeda in the European badger, Meles meles, extending the known host range for this parasite in Europe.
... The temporal activity patterns of a species are affected by physiological processes and the ecological interaction of an Fig. 3 Patterns of daily activities of dhole-banteng, dhole-buffalo, dhole-deer, and dhole-muntjac animal with the environment, as well as biotic factors (Castro-Arellano and Lacher 2009;Zalewski 2001). It is an outcome of conflicting goals, such as maximization of nutritional intake and reproductive success, and the demand to reduce costs and minimize predation risks (Vieira et al. 2017). ...
An understanding of the interspecific interactions within communities is required to explain the mechanisms promoting carnivore co-existence and how prey respond behaviorally to predators. In this research, inter-predator and predator–prey interactions in the tropical savanna and deciduous forests in Baluran National Park, East Java, Indonesia, were investigated. The pattern of interactions was assessed with respect to intraguild predation theory for inter-predator interactions and landscape of fear theory for predator–prey interactions. Data from camera traps were used to examine spatiotemporal activities of animals. The results indicated no evidence of spatial avoidance of solitary predator leopards (Panthera pardus ssp. melas) caused by the presence of social predator dholes (Cuon alpinus), and the data showed that the probability of the occupancy of leopards increases when dholes are present. Within the predator–prey interaction, only dhole and banteng (Bos javanicus) showed a significantly negative spatial interaction. The probability of occupancy of Javan deer increased when dholes and leopards were present; that of the muntjac and the buffalo increased when dholes were present and decreased when leopards were present. By contrast, the segregation of temporal activity was confirmed for all pairs of species, both for inter-predator and predator–prey species. The findings of this research affirmed that temporal interaction was stronger than spatial interaction in determining the inter-predator and predator–prey co-occurrence in the tropical savannah and deciduous forests.
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We review den and resting site terminology used in 121 scientific publications related to species of the subfamily Guloninae (hereafter Martes Complex, sensu Proulx and Aubry 2017). These indicate that the term den has been used both to describe structures used by females to give birth and/or raise their kits, or by both sexes to sleep or rest. The term resting site has been used to describe locations where animals are inactive for varying durations, but no unilateral and systematic link with the den terminology has been established. In the absence of unambiguous, explicit definitions in 53-65% of papers, the lack of consensus on the meaning of these terms hampers efforts towards intra-and interspecific comparisons and sound management plans. Based on the literature and current knowledge on the biology and spatial ecology of species of the Martes Complex, we propose a series of unequivocal definitions for resting (rest) site, den, reproductive den, natal den, and maternal den, and show how these terms are interrelated. We recommend that these definitions be used in future studies, or else that researchers define explicitly these terms in papers dealing with the resting ecology of this group of mesocarnivores.
Red squirrel (Sciurus vulgaris) range within the United Kingdom (UK) has retracted significantly due to the spread of an Invasive Alien Species, the North American Eastern grey squirrel (Sciurus carolinensis). Where grey squirrels are sympatric, red squirrel populations decline through inter-specific competition and squirrelpox virus (SQPV) infection. Grey squirrel eradication from the island of Anglesey facilitated the complete restoration of native red squirrels. Although native species recovery delivered significant ecological and economic benefits, the eradication extended only to a narrow sea-channel boundary, across which grey squirrel dispersal continues to occur. Hence, the long-term sustainability of Anglesey's red squirrel population is vulnerable to grey squirrel re-establishment without continuous intervention. Recent research has demonstrated that as pine marten (Martes martes) landscape use intensity increases, so too does red squirrel occupancy, likely linked to parallel declines in grey squirrel occupancy. Restoration of this mustelid predator is a potential tool to deliver sustainable grey squirrel control by restoring a missing trophic component, depressing grey squirrel incursion rates onto Anglesey, reducing red squirrel exposure to SQPV. Recent UK pine marten translocations have sourced animals under licence from wild Scottish populations. We explore the alternative use of captive-bred founders, simultaneously introducing new genetic variability against a limited diversity within extant populations. A current conservation translocation is paired with an ongoing assessment of founder behaviour and ‘personality’ measured before release. We then highlight the multi-disciplinary approach to delivering applied red squirrel conservation programmes in the face of invasive species.
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The breeding time activity of the Pygmy Owl was studied with automatic registering apparatus in Oulu (65 deg. N) in June 1969. The Owl was most active between 22-01 and 9-10 o'clock. The circadian rhythm of the Pygmy Owl is directly influenced by the light.The food was studied in 1969 in Oulu, Kuusamo and Jämsä. In Oulu, the female and three young ate on average 4.6 preys in 24 hours, From 154 preys 57% were Bank and Field Voles and 38% birds. There was variation in the food at three nests in different parts of Finland due the differences in the prey species available. First time the activity of the Pygmy Owl was compared with the activity of the main prey, the Bank Vole. The same activity peaks were found in both the owl and voles.
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The breeding time food of the Pygmy Owl was studied in three nests in Kuusamo, Oulu and Jämsä in Finland. A total of 255 prey animals were identified in 1969. Bank and Field Voles formed 56.1-74.2% of the diet but birds were also commonly taken (22.2-38.3%). Remaining food consisted shrews, lizards, fish and insects. This could be one of the first times when the activity of the owl is compared with the activity of its main prey. The peaks in the activity of the Bank Vole coincided with those of the Pygmy Owl both being mainly nocturnal but having a clear peak also before noon at 9-10 o'clock. Also the Field Voles are in summer most active in the night when the Pygmy Owl is hunting. Small birds preyed on by the Pygmy Owl are mostly diurnal species.
This book is about the relationships between the entire community of predators and populations of their prey in the Białowieża Primeval Forest. Although the ecological term ‘predator’ is applicable to taxonomically diverse animals, in this work it is used to mean mammals in the order Carnivora and birds of prey in the orders Strigiformes, Accipitriformes, and Falconiformes. We have studied this unique web of relationships for over a decade (1985–1996). Since Białowieża is among the best preserved but also the most thoroughly studied places in the world, we enjoyed the opportunity of combining our data with the results of numerous botanical, zoological, and ecological studies conducted there during the recent half-century. Furthermore, we exploited the wealth of information from the archives of forestry and game management of the forest.
The authors’ intention is to present some selected problems of vegetation and plant population dynamics based on research carried out for many years in the Białowieża Primeval Forest and to confront them in some cases with parallel investigations performed in other natural environments of North-Eastern Poland.
The significance of biological rhythms can be discussed under at least two aspects. They serve, on the one hand, to attain an optimal temporal arrangement of animal behaviour within the cycles of the environment, as in the four “circa-clocks” (Aschoff 1981). On the other hand, this external adaptation results in internal temporal order which in itself may have selective value. In addition, there are many rhythmic processes within the organism, not related to any environmental periodicity, which in various ways contribute to the maintenance of functional integrity of the internal milieu (Aschoff and Wever 1961). In focussing on how circadian rhythms contribute to survival, we do well to consider them, first, as part of a spectrum of rhythms and to evaluate their possible intrinsic function regardless of the environmental day-night cycle. We then will proceed to a discussion of possible benefits to be derived from the adjustment to the periodic environment.