25
Grazing, abandonment and frequent 
mowing influence the persistence of the 
steppe marmot, Marmota bobak
Abstract
The goals of the study were to illustrate how parameters of steppe marmot set-
tlements change after abandonment of livestock grazing, to evaluate differences 
in vegetation characteristics between grazed and abandoned habitats, to find the 
links between these characteristics and the persistence of marmot settlements 
and to establish whether it is possible to maintain the food base of marmots by 
applying frequently repeated mowing. Data were collected in 1991–2016 in the 
steppe marmot (Marmota bobak) settlement in the Regional Landscape Park “The 
Velykyi Burluk-Steppe” (Kharkiv Region, Northeastern Ukraine). We found that 
grazing prolongs the period of active plant growth and supports a regrowth during 
periods of intensive feeding of all age groups of marmots. In abandoned habitats 
the vegetation period ended before the end of active feeding of reproductive fe-
males and juveniles. This resulted in lower reproductive success, which determines 
a decrease in the population and may in the future result in a larger decline of set-
tlements. Frequently repeated mowing by a lawnmower at a cutting height of 6–9 
cm prolongs the vegetation period and maintains a water content in the plants 
similar to that of grazed habitats throughout the active season of steppe marmots. 
Izvleček
Cilj raziskave je bil ponazoriti kako se dejavniki naselbine/kolonije stepskega 
svizca spremenijo po opustitvi paše, ovrednotiti razlike v lastnostih vegetacije 
med pašenimi in opuščenimi habitati, najti povezavo med temi lastnostmi in 
obstojnostjo naselbin/kolonij stepskega svizca in ugotoviti ali je moč ohranjati 
osnovno hrano za svizce z pogosto košnjo. Podatke smo zbrali med letoma 1991 
in 2016 v naselbini/koloniji stepskega svizca (Marmota bobak) v Regionalnem 
parku “Stepa Velykyi Burluk” (regija Kharkiv, severovzhodna Ukrajina). Ugotovili 
smo, da paša podaljša aktivno rast rastlin in omogoča ponovno rast v obdobju 
hranjenja svizcev vseh starostnih skupin. Na opuščenih habitatih se vegetacijska 
doba zaključi pred obdobjem hranjenja plodnih samic in mladičev. To se odrazi v 
zmanjšanem reproduktivnem uspehu in zmanjšanju populacije ter v prihodnosti 
tudi v večjem zmanjšanju števila naselbin/kolonij. Pogosta košnja s kosilnico na 
višini 6–9 cm podaljša vegetacijsko dobo in ohranja vsebost vlage v rastlinah, 
podobno kot paša stepskih svizcev med aktivno sezono.
Keywords: fattening, growing 
season, regrowth, rodent, water 
content. 
Ključne besede: debeljenje, rastna 
sezona, ponovna rast, glodalci, 
vsebnost vlage.
Received: 20. 2. 2017
Revision received: 30. 6. 2017
Accepted: 12. 10. 2017
Co-ordinating Editor:  
Orsolya Valkó
1 Department of Zoology and Ecology, V.N.Karazin Kharkiv National University, Svobody sq. 4, Kharkiv, 61022, Ukraine; E-mail: savchgala5@gmail.com.
Galina Savchenko1 & Vladimir Ronkin1
DOI: 10.1515/hacq-2017-000917/1 • 2018, 25–34
17/1 • 2018, 25–34
26
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
Introduction
Livestock grazing is considered to have an important 
anthropogenic impact on grassland ecosystems with 
a variety of effects (Dengler et al. 2014). In order to 
mitigate the negative effects of overgrazing, which 
remains a problem for many countries, sustainable pasture 
management strategies are proposed to be implemented 
(Baranova et al. 2016). Finding the best grazing practice 
is proposed as a solution for many problems of Palearctic 
grasslands, such as maintaining the optimal level of bio-
diversity, suppression of weed growth (Török et al. 2014, 
2016) and support of steppe-like vegetation and arthro-
pod assemblages (Polchaninova et al. 2016). Abandon-
ment of grazing has been an object of studies as well. It 
was shown that pasture abandonment leads to an increase 
in vegetation height and in the richness of mesophytes, 
plants with leafy canopy and red-list species, whereas 
grazed plots are often characterised by higher total spe-
cies richness and more xerophytes, rosette forming and 
spring-flowering species (Vassilev et al. 2011). 
Conservation management strategies for grasslands 
should be based on the knowledge of local ecological 
specifics of these sensitive systems (Valkó et al. 2016). 
The results of various studies of maximum number of 
their components including herbivores and granivores 
mammals should be taken into account as well. Previous 
studies reported that intensive grazing negatively affects 
the abundance and diversity of small mammals (Yarnell 
et al. 2007). Effects of management actions on grassland 
mammals may be mediated by vegetation height (Mérő 
et al. 2015), quality of available food (Abaturov 2006) 
and proportion of food plants in the total aboveground 
phytomass (Ronkin & Savchenko 2000).
Grazing in the grasslands of Northeastern Ukraine may 
be completely abandoned in the nearest future due to signif-
icant changes in livestock farming. One of the endangered 
open landscape rodents, the steppe marmot (Marmota bo-
bak), is a secondary user of these grazed grasslands (Ronkin 
et al. 2009). Similar to the Alpine marmot (Arnold 1990, 
Perrin et al. 1993, Lenti Boero 1996, 2001), all individuals 
of a family of steppe marmots hibernate together in the 
same winter burrow. During vegetation season marmots 
must obtain the required amount of food for reproduction, 
growth and fat storage for hibernation (Frase & Armitage 
1989, Bibikov 1989, Melcher et al. 1989, Mashkin 1997). 
These rodents consume a large number of plant species 
(Chovankova & Soltesova 1988, Bassano et al. 1996, Her-
rero et al. 2002), but they do not feed in proportion to the 
species abundance in the environment; instead, a few food 
species may fulfil a large part of their nutritional needs 
(Massemin et al. 1996, Ronkin & Savchenko 2004).
The goals of the study were: a) to illustrate how param-
eters of the marmot settlements change after abandonment 
of cattle grazing, b) to evaluate differences in vegetation 
characteristics of grazed and abandoned habitats that are 
important for the marmots’ food base, c) to find the links 
between the habitat characteristics and the persistence of 
marmot settlements and d) to establish whether it is pos-
sible to maintain characteristics of the vegetation similar to 
grazed habitats by applying frequently repeated mowing.
Materials and methods
The study was conducted in 1991–2016 in the steppe 
marmot settlement in the Regional Landscape Park “The 
Velykyi Burluk-Steppe” (Kharkiv Region, Northeastern 
Ukraine). In this part of the Kharkiv Region, the mean 
annual rainfall is about 540 mm with a peak in June 
(63 mm), and the mean annual temperature is 7.0 °C, 
with July and August temperatures often exceeding 36 °C. 
In January the mean temperature ranges from −6.8 °C to 
−8.5 °C, with a minimum temperature of −35 °C.
The permanent observation plot was located near the 
village of Nesterivka (49° 55' 19" N; 37° 18' 46" E). The 
studied area of the marmot habitats is about 85 ha in size. 
The land use history, relief, common management prac-
tices, flora and vegetation of this permanent study area 
were described in detail in a previous publication (Ronkin 
& Savchenko 2016).
Sampling methods
The number of marmot families and the density of the 
marmot population were estimated by counting the 
winter burrows of a given territory. For the mapping of 
inhabited and uninhabited winter burrows, a handheld 
Global Positioning System device (Garmin Oregon 600 
Series) and the program QGIS v.2.18.8 were used.
Observations were made in activity hours of marmots 
(in spring and autumn during the day, in summer in the 
morning and/or evening); the fieldwork lasted no less 
than 100 hours per year. The following parameters were 
recorded: habitability of a winter burrow, number of fam-
ily members, phase of active season of marmots (emerg-
ing from hibernation, beginning and end of active feed-
ing, juveniles’ first appearance above the ground).
The number of marmots in the family was counted 
with binoculars and spotting scopes when the animals 
were feeding freely. Individuals were counted and grouped 
into the following three categories: adult males + non-re-
productive females, reproductive females (females having 
offspring in the current year) and juveniles. Whether an 
17/1 • 2018, 25–34
27
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
observed animal belonged to a particular family was de-
termined by its position. This approach was chosen since 
steppe marmot family members use a certain territory that 
does not overlap with home ranges of other families. The 
size of their home range varies in different habitats from 
about 0.6 to near 2.5 ha (Mashkin 1997). At least one 
winter burrow, foraging areas and refuges are the obliga-
tory elements of their home range. This spatial structure of 
the settlement has been determined in previous studies on 
tagged marmots (Nikol’skii & Savchenko 1999). Because 
the positions of the obligatory elements of a home range 
do not change for years and borders of families are stable 
(Mashkin 1997), the family membership of an observed 
animal may be distinguished on the basis of its position.
Observations of marmot behavior and appearance 
helped to determine the beginning and the end of certain 
phases of the marmots’ active season. In this research we 
were interested in the periods of active feeding (fattening 
period). The predominance of feeding activity marks the 
start of the fattening period, and completion of shedding 
and accomplishment of fatness marks its end (Bibikov 
1989, Mashkin 1997).
Vegetation characteristics and 
management regimes
The grasslands of Northeastern Ukraine were formed by 
livestock grazing (mainly cattle and sheep). In the 1990s 
maximal grazing pressure was 4–5 cows/ha. Because “The 
Velykyi Burluk-Steppe” is a gully net (Figure 1), the vege-
tation of the marmot habitats is characterised by commu-
nities of both the forb-fescue-feather grass steppe (mainly 
Figure 1: Gully relief of the Regional Landscape Park “The Velykyi Burluk-Steppe” (A), heavily grazed habitat of Marmota bobak (1996) (B) and 
habitat after abandonment (C). Photos: Vladimir Ronkin.
Slika 1: Razgiban jarkast relief v regionalnem krajinskem parku “Stepa Velykyi Burluk“ (A), močno popašen habitat vrste Marmota bobak (1996) 
(B) in habitat po opustitvi gospodarjenja (C). Fotografije: Vladimir Ronkin.
17/1 • 2018, 25–34
28
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
Festuca valesiaca agg. and forbs) and steppe meadows 
(mainly Poa angustifolia and forbs) (see also Ronkin & 
Savchenko 2016). Further in the text, these communities 
are referred to as steppe and steppe meadow, respectively.
The following characteristics of the habitats were re-
corded between 2009 and 2016: presence or absence of 
cattle grazing, its intensity, height of the basic grass layer 
and phase of vegetation (active growth, maturing of grass-
es, vegetative rest, senescence).
To evaluate fluctuations in the water content of the veg-
etation in foraging areas of different habitats (grazed and 
abandoned steppes and steppe meadows), vegetation sam-
ples were collected twelve times during the seasons of mar-
mot activity in 2012/2013 (between mid-April and mid-
October). The vegetation was clipped at a height of about 
6 cm and immediately weighed, then dried to a constant 
weight and weighed again. We used a lawnmower for a 
mass collection of the vegetation samples and recommend 
it as a very useful tool for collecting such samples.
Experiments on frequently repeated mowing of the 
vegetation of the habitats after abandonment were carried 
out in 2014/2015. Two areas of 0.01 ha each (a steppe 
and a steppe meadow) were mown by a lawn mower at a 
height of 6 cm during the growing season. A grass area of 
2 m2 was mown to determine the productivity. The fre-
quency of mowing – 20 times in 2014 and 15 times in 
2015 – was conditioned by the regrowth of the basic level 
of the grass up to 8–9 cm. With this very frequent mow-
ing, we wanted to model the effects of heavy grazing. The 
water content was determined in each sample.
Statistical analysis
The average number of individuals of three groups (adult 
males + non-reproductive females, reproductive females 
and juveniles) in a marmot family was estimated on an-
nual basis between 2005 and 2013 and used in a factorial 
ANOVA. As factors, two parameters of the settlement were 
included: factor “habitat” with two levels (steppe or steppe 
meadow), factor “management” with two levels (grazing or 
abandonment) and their combination (habitat × manage-
ment). Since the average number of reproductive females 
and juveniles on the sites after abandonment differed sig-
nificantly from normal distribution (P < 0.05, Shapiro-
Wilk and Kolmogorov-Smirnov normality tests), the U-test 
(Mann-Whitney) was additionally applied. In order to esti-
mate differences in reproduction success on the sites, only 
juveniles that subsequently survived the first hibernation 
were considered in the tests. This approach, i.e. focusing on 
the number of marmots that survived at least one hiberna-
tion, is also used by other marmot specialists, for example 
when analysing group size (Blumstein & Foggin 1997).
The combination of habitat and management allowed 
us to distinguish four sites (“grazed steppe”, “grazed 
steppe meadow”, “steppe after abandonment” and “steppe 
meadow after abandonment”); these names are used fur-
ther in the text. The Mann-Whitney test was used in the 
water content analysis (where “habitat”, “management” 
and “year” were included as factors). For each site, the 
durations of the growing season in 2009–2016 were ana-
lysed using Kruskal-Wallis ANOVA by ranks, and the end 
dates of the fattening period of different marmot groups 
together with the dates of the onset of vegetation rest 
were analysed using one-way ANOVA (the variables were 
tested for normality).
Statistical significance was set at P < 0.05. The Least 
Significant Difference (LSD) criterion was used for means 
comparison relating to phases (periods) in plants and 
marmots. All calculations were performed in the program 
StatSoft Statistica v6.0.
Results
Characteristics of the marmot 
settlement
In 1991, 51 marmot families were registered in the settle-
ment under study. At that time all marmot habitats were 
heavily grazed by cattle. The grazed area began to decrease 
in 1992, and by 2014, only 10 marmot families stayed in 
the ungrazed part of the settlement, which was 36% of 
the families compared to 1992. The population density 
decreased from 1.2 to 0.4 families/ha on the site “steppe 
meadow after abandonment” and from 0.8 to 0.2 fam-
ilies/ha on the “steppe after abandonment”. In contrast, 
the number of families within the zone of intensive graz-
ing remained on the same level as it was in 1992 (23 fam-
ilies) (Figure 2).
We found that only one factor – “management” – had 
a statistically significant influence on the number of indi-
viduals in a marmot family (factorial ANOVA, F = 16.9, 
df = 3, 30, P < 0.001) (Figure 3). The average number of 
individuals was lower in both habitats “after abandon-
ment” for all the groups of marmots, especially for repro-
ductive females (U = 36.0, n1 = n2 = 18, P < 0.0001) and 
juveniles (U = 21.0, n1 = n2 = 18, P < 0.0001). On the 
site “steppe after abandonment”, an appearance of juven-
iles was registered only once during 2005–2013, but no 
young animals emerged after the first hibernation. On the 
site “steppe meadow after abandonment”, an appearance 
of offspring was observed three times. The total number 
of juveniles was ten, and only four of them emerged after 
the first hibernation.
17/1 • 2018, 25–34
29
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
15
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grazed steppe meadow
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adult males and non-
-reproductive females
Vegetation characteristics and 
management actions
The highest grass was registered in the “steppe meadow 
after abandonment”. The basic level was at its highest 
from mid-June to late July (mean±SD: 64.6±7.4 cm) 
(Figure 4). In the “steppe after abandonment”, the same 
characteristic gained its maximum from the middle of 
May to the end of June (mean 23.7±4.5 cm). On the 
grazed areas, the height was 4–7 cm during the whole 
grazing season (from early May to late September/early 
October).
Figure 3: Mean number of individuals in a marmot family on grazed 
and abandoned sites (factorial ANOVA, F = 16.91, df = 3, 30, 
P <0.001).
Slika 3: Povprečno število osebkov v družinski skupnosti svizcev na 
pašenih in opuščenih habitatih (faktorska analiza variance ANOVA, 
F = 16,91, df = 3, 30, P <0,001).
Figure 2: Location of marmot families in grazed and abandoned parts 
of the settlement in the Regional Landscape Park “The Velykyi Burluk-
Steppe” near the village of Nesterivka (2014).
Slika 2: Lokacije kolonij svizcev v pašenem in opuščenem delu nasel-
bine v regionalnem krajinskem parku “Stepa Velykyi Burluk“ pri vasi 
Nesterivka (2014).
Figure 4: Height of the basic grass layer on different sites (mean of 
averages for the years 2012–2015).
Slika 4: Višina osnovne travniške plasti na različnih rastiščih (povprečja 
za leta 2012–2015).
Fluctuations of water content in the vegetation in the 
communities showed similar trends in 2012 and 2013 
(Figure 5). The vegetation of the “grazed steppe meadow” 
was not only the wettest (with a mean water content of 
72.3±4.4% in 2012 and 73.4±2.9% in 2013), but also 
that with the smallest fluctuations throughout the grow-
ing season. The vegetation of the “steppe after abandon-
ment” showed the opposite characteristics (with a mean 
water content of 45.3±19.2% and 39.7±18.6%, respec-
tively). The water content was statistically significantly 
dependent on “management” (U = 88.7, n1 = n2 = 44, 
P < 0.001). The effects of “habitat” (U = 784.5, n1 = 
n2 = 44, P = 0.13),“year” (U = 862.5, n1 = n2 = 44, 
P = 0.38) and all interactions of the factors were not sta-
tistically significant.
On the sites where frequently repeated mowing was 
conducted, the grass was kept at a height of 6–9 cm dur-
ing the whole growing season. In May of both years (Fig-
ure 6), the regrowth in the steppe was larger than in the 
steppe meadow, especially in 2015, when the weather was 
unusually warm and rainy. The total regrowth for the en-
17/1 • 2018, 25–34
30
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
tire growing season in 2014 amounted to 397.13 g/m2 for 
the steppe meadow and 355.2 g/m2 for the steppe (dry 
weight); in 2015, the figures were 227.05 and 305.5 g m2, 
respectively. As for the average daily growth (Figure 6), 
both communities had similar figures (except in May 
2015) and similar fluctuations during the growing season. 
The water content on the mown sites (Figure 7) was 
similar to that of the grazed ones (Figure 5). Almost 
throughout the season, except the first half of May, the 
water content was slightly higher in the steppe mead-
ow: on the grazed sites, the mean values in 2012/2013 
were 72.8±3.7 vs. 68.4±6.3%; on the mown sites in 
2014/2015, they were 70.6±5.1 vs. 68.1±6.9%.
We recorded that the duration of the growing season 
in 2009–2016 ranged from 189 to 231 days. The longest 
period of active plant growth was registered at the site 
“grazed steppe meadow” and the shortest at the “steppe 
after abandonment” (219±14 vs. 101±9 days, n=8). The 
growing season on the sites “grazed steppe” and “steppe 
meadow after abandonment” was usually interrupted by 
a drought in the second half of the summer (Figure 8); 
this happened seven times between 2009 and 2016. 
The completion of the growing season on the sites 
where frequent mowing was conducted occurred at 
approximately the same time as in the respective grazed 
communities.
Growing season and fattening 
period of marmots
As shown in Figure 8, the period of intensive feeding of 
animals started after the onset of the growing season; it 
was shortest for adult males and non-reproductive fe-
Figure 5: Water content of plant biomass during the growing seasons of 2012 (A) and 2013 (B) in the grazed and abandoned communities.
Slika 5: Vsebnost vode v rastlinski biomasi med rastno sezono 2012 (A) in 2013 (B) v pašenih in opuščenih združbah.
Figure 6: Daily plant regrowth during the growing seasons of 2014 
and 2015 in the communities with frequently repeated mowing.
Slika 6: Dnevni prirastek med rastnima sezonama 2014 in 2015 v 
združbah s pogosto košnjo.
Figure 7: Water content of the plant biomass during the growing 
seasons of 2014 and 2015 in the communities with frequently repeated 
mowing.
Slika 7: Vsebnost vode v rastlinski biomasi med rastnima sezonama 
2014 in 2015 v združbah s pogosto košnjo.
grazed steppe grazed steppe meadow steppe after abandonment steppe meadow after abandonment
23
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2015 steppe
2015 steppe meadow
2014 steppe
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2014 steppe 2014 steppe meadow
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2015 steppe meadow
2014 steppe
2014 steppe meadow
w
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17/1 • 2018, 25–34
31
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
males and longest for juveniles. Besides, adult males and 
non-reproductive females began the intensive feeding 
earlier than reproductive females and juveniles (the differ-
ence was 20±7 days).
The average duration of the period of active plant growth 
differed significantly between all the sites (Kruskal-Wallis 
test, H = 25.5, df = 3, P < 0.001). The differences in the 
average end dates of the active feeding period of different 
marmot groups as well as in the onset of the vegetation 
rest at the different sites were also statistically significant 
(one-way ANOVA, F = 80.9, df = 6, 49, P < 0.001). The 
“grazed steppe meadow” was characterised by the longest 
period of active plant growth, which covered the periods 
of intensive feeding of three groups of marmots (LSD0.05 = 
12.45, P < 0.001). In both communities “after abandon-
ment”, such covering did neither occur for reproductive 
females nor for juveniles (LSD0.05 = 12.45, P < 0.01). In 
the “grazed steppe”, the covering was recorded six times 
in eight years for reproductive females and four times for 
juveniles.
Discussion
According to estimates of marmot specialists, the above-
ground phytomass in habitats of these rodents ranges 
from about 22 g/m2 (in case of golden marmots, Blum-
stein & Foggin 1997) to about 800 g/m2 (in case of 
yellow-bellied marmots, Frase & Armitage 1989). It was 
shown that in poor habitats (average total aboveground 
productivity from 21.9 g/m2 to 51.1 g/m2) of high alpine 
meadows, the probability of weaning juveniles was as-
sociated with the overall food ability of the previous 
year, and there was a significant positive relationship 
between the early season food resources and the propor-
tion of years in which a group reproduced (Blumstein & 
Foggin 1997). The estimated amounts of total regrowth 
obtained by us for the entire growing season ranges be-
tween 200 g/m2 and 400 g/m2. The maximum and mini-
mum values of the average daily regrowth in the studied 
communities as well as the difference in dynamics be-
tween 2014 and 2015 may be explained by differences in 
weather conditions. Obviously, the mass of food in the 
steppe marmot habitats is enough to meet the needs of 
the animals as it is considered by other authors (Mashkin 
1997). In this research we focus not so much on the total 
mass of food, but on its quality, which may be expressed 
by water content. It was observed in experiments (Ron-
kin & Tokarsky 1993) that marmots selected younger 
and thus juicier parts of plants with water content of no 
less than 70%. If converted to a dry weight of the food, 
the animals selected parts, which contained more pro-
tein (16.0 vs. 9.2%), nitrogen-free extract (NFE) (56 vs. 
46%), and energy (1586.4 vs. 1418.1 kJ/100 g) than the 
rejected parts. Considering the dynamics of the water 
content, the “grazed steppe meadow” is the most favour-
able. In contrast, the water content in the vegetation on 
both sites “after abandonment” decreases rapidly, i.e., the 
vegetation ages, and its nutritional quality drops.
The degree of correspondence between the growing 
season of food species and the fattening period of mar-
mots with different reproductive status and age essentially 
affects reproduction in marmots (Bibikov 1989). It was 
found that precipitation, length of growing season and 
period of active growth of plants
period of temporary stopping of plants growth
fattening period in adult marmots and period of growth in juveniles
Feb Mar Apr May Jun Jul Aug Sep Oct Nov
steppe after abandonment
grazed steppe
grazed steppe meadow
steppe meadow after abandonment
juvenile marmots
reproductive females
adult males and non-reproductive females
Figure 8: Mean terms and lengths (2009–2016) of the fattening period of marmots with different reproductive status and age in comparison with 
the growing season (from start of active growth to senescence of vegetation) in the grazed and abandoned habitats.
Slika 8: Povprečen začetek in trajanje obdobja (2009-2016) debeljenja svizcev z različnim reproduktivnim statusom in starostjo v primerjavi z 
rastno sezono vegetacije (od začetka do prenehanja rasti vegetacije) na pašenih in opuščenih habitatih.
17/1 • 2018, 25–34
32
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
length of winter were the major factors affecting reproduc-
tion and survival of yellow-bellied marmots in the Upper 
East River Valley, Colorado (USA) (Schwartz & Armitage 
2002, Armitage 2013). On a basis of energy allocation 
by yellow-bellied marmots, Melcher et al. (1989) showed 
that a necessary minimum fat deposit was crucial for the 
survival of juveniles during hibernation and that the total 
energy demand was highest for reproductive females. In 
our study a longer growing season in grazed habitats pro-
vided periods of foraging activity for juveniles and repro-
ductive females that allowed them to continue foraging 
when vegetation in abandoned habitats was already un-
suitable as food. The “steppe after abandonment” becomes 
mature in June, and their growing season corresponds to 
the fattening period of adult males and non-reproductive 
females only. The “steppe meadow after abandonment” 
has a longer growing season, but the vegetation period is 
interrupted in the summer. To survive the first hiberna-
tion, juveniles need a longer and uninterrupted growing 
season. Cattle grazing prolongs the growing season, which 
is extremely beneficial for the survival of juveniles.
In general, grazing affects other characteristics of the 
food base, too. First of all, it has an impact on the spe-
cies composition. Forbs, which are common in heavily 
grazed areas, are the basic food species for marmots, i.e. 
the plants eaten by these rodents during all the active 
season (Ronkin & Savchenko 2000). Another important 
characteristic is the ratio between the palatable and the 
non-palatable part of the vegetation. A simultaneous in-
crease in grass height and decrease in water content means 
a decrease in the proportion of palatable vegetation on the 
sites “after abandonment”. In this case marmots experi-
ence a food deficiency starting in mid-June, i.e. before the 
end of the growing season.
Probably, this lack of food is the main cause of a decline 
in the number of reproductive females and an even more 
drastic decrease in the number of surviving juveniles. Due 
to the relatively long lifespan of steppe marmots, the pro-
cess of settlement depletion after abandonment of grazing 
is relatively slow. In addition, this process is slowed down 
by migration (as a result of the lack of free home ranges) 
of marmots from favorable parts of the settlement. Even 
though, the abandonment of grazing in the entire terri-
tory of a settlement will inevitably lead to a complete de-
population of those settlements in 5–7 years. This term is 
determined by the fact that the proportion of reproduc-
tive females declines to 5% as they reach the age of 5 years 
(Mashkin 1997).
The most successful part of the settlement occupies 
heavily grazed habitats with mainly mesic non-senescent 
vegetation of low height, which has a prolonged vegeta-
tion season and constant regrowth as well as an abun-
dance of food species. Th e “grazed steppe” does not 
support the completion of the fattening in reproductive 
females and juveniles every year: Two times in eight years, 
the second half of both the growth period of juveniles and 
the fattening of reproductive females were interrupted by 
a temporary interruption of plant growth. However, in 
grazed part of the settlement, most of the marmots’ home 
ranges include both communities. On the one hand, this 
provides the fattening of all the groups of marmots, and 
on the other hand, it explains the prevailing influence of 
the “management” factor on the number of individuals in 
marmot families. Like grazing, frequently repeated mow-
ing reduces the height of the grass, promotes regrowth 
and prolongs the active growing season. The duration of 
the growing season is similar to that of grazed sites and 
covers the fattening periods of all groups of marmots.
Obviously, on a larger scale (of a whole settlement or 
population), only grazing can maintain vegetation char-
acteristics necessary for the reproduction of steppe mar-
mot populations. However, frequently repeated mowing 
can serve as a substitute for intensive grazing at a smaller 
scale comprising at least the most favorable parts of a set-
tlement. Taking into account the tendencies in the devel-
opment of livestock farming, it is easier to realise such a 
frequent mowing than to reintroduce the former grazing 
regimes. The use of this practice allows maintaining the 
vegetation characteristics necessary for the steppe marmot 
reproduction. The patchy pattern of mowing will not only 
support the marmots’ nutritional needs, but also take 
into account the requirements of other grassland dwell-
ers, which are reported to respond negatively to mowing 
(Polchaninova 2004, Cizek et al. 2012, Mazalová et al. 
2015). This approach supports the idea of zoning practice 
(Vassilev et al. 2011) and promotes the application of a 
practice that is heterogeneous in space and time (Török 
et al. 2016).
Conclusions
Grazing prolongs the period of active plant growth and 
supports a regrowth during periods of intensive feeding of 
all the age groups of marmots. Abandonment of grazing 
negatively influences the persistence of the steppe marmot 
settlements. In abandoned habitats, the vegetation period 
ends before the end of the active feeding of reproductive 
females and juveniles. This is the cause of reduction in re-
productive success, which determines the decrease of the 
population density and may result in a complete depopu-
lation of settlements in the future. At a larger scale, only 
grazing can maintain vegetation characteristics neces-
sary for the reproduction of steppe marmot populations. 
17/1 • 2018, 25–34
33
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
However, frequenty repeated mowing with a lawnmower 
at a cutting height of 6–9 cm prolongs the vegetation pe-
riod and maintains a water content in the plants similar 
to that of grazed habitats throughout the active season of 
steppe marmots. We recommend this practice to save the 
marmots’ food base in the most favorable parts of marmot 
settlements. In general, considering not only the needs 
of marmots, but also other species of flora and fauna, 
we suggest that spatially and temporally heterogeneous 
(patchy) management practices may be useful.
Acknowledgments
The authors would like to thank Orsolya Valkó as the 
Co-ordinating Editor. The authors are grateful to two 
anonymous reviewers and Orsolya Valkó for the editing 
the manuscript and for the recommendations, comments 
and improvements of former drafts of the paper. We 
also thank the family of Alex Zalyvadniy (Vancouver, 
Canada) for the improvement of the English text. We 
would like to express our sincere gratitude to Dr. Aiko 
Huckauf for the professional linguistic editing, and the 
Eurasian Dry Grassland Group (EDGG) for possibility 
to publish this paper.
References 
Abaturov, B. D. 2006: Pastbishchniy tip funktsionirovaniya stepnykh 
i pustynnykh ekosistem (Grazing type of functioning of steppe and 
desert ecosystems). Ouspekhi Sovremennoy Biologii 126(5): 435–447. 
[In Russian].
Armitage, K. B. 2013: Climate change and the conservation of 
marmots. Natural Science 5(5A): 36–43.
Arnold, W. 1990: The evolution of marmot sociality: I. Why disperse 
late? II. Costs and benefits of joint hibernation. Behavioral Ecology 
and Sociobiology 27: 229–246.
Baranova, A., Schickhoff, U., Wang, S. & Jin, M. 2016: Mountain 
pastures of Qilian Shan: plant communities, grazing impact and 
degradation status (Gansu province, NW China). Hacquetia 15(2): 
21–36.
Bassano, B., Peracino, V., Pecacino, V. & Montacchini, F. 1996: Diet 
composition and feeding habitats in a family group of alpine marmot 
(Marmota marmota) – preliminary data. In: Le Berre, M., Ramousse, 
R. & Le Guelte, L. (eds.): Biodiversity in Marmots. International 
marmot Network, Moscow, Lyon, pp. 135–140.
Bibikov, D. I. 1989: Surki (The Marmots). Agroprom Press, Moscow, 
255 pp. [In Russian].
Blumstein, D. T. & Foggin, J. M. 1997: Effect of vegetative variation 
on weaning success, overwinter survival, and social group density 
in golden marmots (Marmota caudata aurea). Journal of Zoology, 
London 243: 57–69.
Chovankova, B. & Soltesova, A. 1988: Trophic base and nutritional 
activity of the alpine marmot (Marmota marmota latirostris Kratochvil, 
1961). Zbornic prac o TANAP-e 28: 71–135.
Cizek, O., Zamecnik, J., Tropek, R., Kocarek, P., & Konvicka, M. 
2012: Diversification of mowing regime increases arthropods diversity 
in species-poor cultural hay meadows. Journal of Insect Conservation 
16(2): 215–226.
Dengler, J., Janišová, M., Török, P. & Wellstein, C. 2014: Biodiversity 
of Palaearctic grasslands: a synthesis. Agriculture, Ecosystems & 
Environment 182: 1–14.
Frase, B. A. & Armitage, K. B. 1989: Yellow-bellied marmots are 
generalist herbivores. Ethology Ecology & Evolution 1: 353–366.
Herrero, J., Garsia-Gonzalez, R. & Garcia-Serrano, A. 2002: Research 
on the alpine marmot (Marmota marmota) in the Spanish Pyrenees. In: 
Armitage, K.B. & Rumiantsev, V. Yu. (eds.): Holarctic marmots as a 
factor of biodiversity. ABF P. H., Moscow, pp. 190–197.
Lenti Boero, D. 1996: Space and resource use in Alpine marmots 
(Marmota marmota L.). In: Le Berre, M., Ramousse, R. & Le Guelte, 
L. (eds.): Biodiversity in Marmots. International Marmot Network, 
Moscow, Lyon, pp. 175–180.
Lenti Boero, D. 2001: Occupation of hibernacula, seasonal activity, 
and body size in a high altitude colony of Alpine marmots (Marmota 
marmota). Ethology Ecology & Evolution 13: 209–223.
Mashkin, V. I. 1997: Yevropeyskiy baybak: ekologiya, sokhraneniye i 
ispol’zovaniye (The European bobak: Ecology, Conservation and Use). 
Kirov Regional Press, Kirov, 160 pp. [In Russian].
Massemin, S., Gibault, C., Ramousse, R. & Butet, A. 1996: First data 
on the alpine marmot (Marmota marmota) diet. Mammalia 60(3): 
351–361.
Mazalová, M., Šipoš, J., Rada, S., Kašák, J., Šarapatka, B., & Kuras, 
T. 2015: Responses of grassland arthropods to various biodiversity-
friendly management practices: Is there a compromise? European 
Journal of Entomology 112(4): 734–746.
Melcher, J. C., Armitage, K. B. & Porter, W. P. 1989: Energy alloca-
tion by yellow-bellied marmots. Physiological Zoology 62: 429–448.
Mérő, T. O., Bocz, R., Polyák, L., Horváth, G. & Lengyel, S. 2015: 
Local habitat management and landscape-scale restoration influence 
small-mammal communities in grasslands. Animal Conservation 18: 
442–450.
Nikol`skii, A. A. & Savchenko, G. A. 1999: Structure of family groups 
and space use by steppe marmots (Marmota bobac): preliminary results. 
Vestnik Zoologii 33(3): 67–72.
Perrin, C., Allainé, D. & Le Berre, M. 1993: Socio-spatial organization 
and activity distribution of the alpine marmot Marmota marmota: 
preliminary results. Ethology 93: 21–30.
Polchaninova, N. Yu. 2004: Effect of hay-mowing on spider 
communities of the meadow steppes of the Central forest-steppe 
(Russia and Ukraine). Arthropoda Selecta. Special Issue No1 
(Proceedings of the 21st European Colloquium of Arachnology, St.-
Petersburg, 4-9 August 2003): 261–273.
Polchaninova, N., Savchenko, G., Drogvalenko, A., Ronkin, V. & 
Shabanov, D. 2016: The impact of cattle grazing on cursorial spiders 
(Aranei) and true bugs (Heteroptera) in steppe gullies of northeastern 
Ukraine. Agriculture, Ecosystems & Environment 234: 65–71.
17/1 • 2018, 25–34
34
G. Savchenko & V. Ronkin
Grazing, abandonment and frequent mowing influence the persistence of the 
steppe marmot, Marmota bobak
Ronkin, V. I. & Savchenko, G. A. 2000: Adaptability of habitats for 
Marmota bobak (Rodentia, Sciuridae) related to plant cover structure. 
Zoologicheskii Zhurnal 79(10): 1229–1234. [In Russian].
Ronkin, V. I. & Savchenko, G. A. 2004: Effect of cattle grazing on 
habitats for the steppe marmot (Marmota bobak) in north-eastern 
Ukraine. Vestnik Zoologii 38(1): 55–60.
Ronkin, V. & Savchenko, G. 2016: Flora and vegetation of dry 
grasslands of Northeastern Ukraine, and problems of diversity 
conservation. Hacquetia 15(2): 49–62.
Ronkin, V., Savchenko, G. & Tokarsky, V. 2009: The place of the 
steppe marmot in steppe ecosystems of Ukraine: An historical 
approach. Ethology Ecology & Evolution 21: 277–284.
Ronkin, V. I. & Tokarsky, V. A. 1993: Qualitative and quantitative 
assessment of feeding habits of bobak, Marmota bobak, and long-tailed 
marmot, M. caudata (Rodentia, Sciuridae) in captivity. Zoologicheskii 
Zhurnal. 72(2): 93–100. [In Russian].
Schwartz, O. A. & Armitage, K. B. 2002: Correlations between 
weather factors and life-history traits of yellow-bellied marmots. In: 
Armitage, K. B. & Rumiantsev, V. Yu. (eds.): Holarctic marmots as a 
factor of biodiversity. ABF P. H., Moscow, pp. 345–357.
Török, P., Valkó, O., Deák, B., Kelemen, A. & Tóthmérész, B. 2014: 
Traditional cattle grazing in a mosaic alkali landscape: Effects on 
grassland biodiversity along a moisture gradient. PLoS ONE 9 (5): 
e97095.
Török, P., Valkó, O., Deák, B., Kelemen, A., Tóth, E. & Tóthmérész, 
B. 2016: Managing for species composition or diversity? Pastoral 
and free grazing systems in alkali steppes. Agriculture, Ecosystems & 
Environment 234: 23–30.
Valkó, O., Zmihorski, M., Biurrun, I., Loos, J., Labadessa, R. & 
Venn, S. 2016: Ecology and Conservation of Steppes and Semi-natural 
Grasslands. Hacquetia 15(2): 5–14.
Vassilev, K., Pedashenko, H., Nikolov, S. C., Apostolova, I. & Dengler, 
J. 2011: Effect of land abandonment on the vegetation of upland 
semi-natural grasslands in the Western Balkan Mts., Bulgaria. Plant 
Biosystems 145: 654–665.
Yarnell, R. W., Scott, D. M., Chimimba C. T. & Metcalfe, D. J. 2007: 
Untangling the roles of fire, grazing and rainfall on small mammal 
communities in grassland ecosystems. Oecologia 154: 387–402.