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Effect of the Number of Mating Partners and Sperm Quality on Reproductive

Success in the Domestic Cat (Felis catus)

Article  in  Biology Bulletin · December 2018

DOI: 10.1134/S1062359018070063

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ISSN 1062-3590, Biology Bulletin, 2018, Vol. 45, No. 7, pp. 756–765. © Pleiades Publishing, Inc., 2018.
Original Russian Text © M.N. Erofeeva, G.S. Alekseeva, P.A. Sorokin, S.V. Naidenko, 2017, published in Zoologicheskii Zhurnal, 2017, Vol. 96, No. 10, pp. 1243–1253.

Effect of the Number of Mating Partners and Sperm Quality

on Reproductive Success in the Domestic Cat (Felis catus)
M. N. Erofeevaa, *, G. S. Alekseevaa, P. A. Sorokina, and S. V. Naidenkoa
aSevertsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, 119071 Russia
*e-mail: erofeevamariya@yandex.ru
Received October 27, 2016

Abstract—Data on the reproductive success and reproductive behavior of the domestic cat, the most wide-
spread representative of the family Felidae, are presented. The effects of the number of partners, mating
order, and sperm quality on male and female reproductive success were assessed. The experiments showed
that the fertility of female cats that mated with two partners was higher. Here, male reproductive success was
less dependent on the attitude of females to the males, whereas the sperm quality (percentage of morpholog-
ically intact spermatozoa) and order of mating with the female had considerable effects on reproductive suc-
cess. However, male sperm quality did not have a significant effect on the number of offspring if the mating
pattern was monogamous.

Keywords: reproductive success, sperm quality, mating pattern, Felis catus

DOI: 10.1134/S1062359018070063

INTRODUCTION connection has been demonstrated in some mamma-

The domestic cat (Felis catus) is the most common
feline species that forms free-living populations of
feral individuals on all continents (except for the Ant-
arctic) and a number of oceanic islands. This species is
largely dependent on the human population, or actu-
ally, on the population density and the degree of hab-
itat transformation by human activity. As a rule, the
feral cat population density is much higher in cities
than in the countryside (2000–3000 individuals per
1 km2 versus 0.25 individual/km2) (Liberg and Sand-
ell, 1988; Pavlova et al., 2015). This variability of the
population density, in turn, brings about considerable
differences in the feral cat lifestyle. The domestic cat is
a mostly solitary and monogamous animal at a low
population density (Liberg, 1981; Pontier and Natoli,
1996; Say et al., 1999, 2002) and a social and promis-
cuous animal at a high population density (Izawa et
al., 1982; Yamane, 1998; Say et al., 1999, 2002).
Teratozoospermia is relatively uncommon in
domestic cat males, in contrast to most species of the
Felidae family (Wildt et al., 1986, 1988; Howard and
Wildt, 1990, 1994; Pukazhenti et al., 2001, 2006;
Goeritz et al., 2006). Teratozoospermia is currently
recognized as a feature of 28 (of 37) species of the fam-
ily, that is, more than half of all males of these species
are characterized by a high (over 60%) share of anom-
alous spermatozoa in the ejaculate (Pukazhenti et al.,
2006). Low genetic diversity is presumably the pri-
mary reason for the high incidence of teratospermia in
felines (Wildt et al., 1983; Wildt, 1994): at least, such a
lian species (Gage et al., 2006; van Eldik et al., 2006;
Asa et al., 2007; Naidenko et al., 2007).
The low incidence of teratospermia in domestic cat
males may be due to the high abundance and broad
living range of this species (Legay, 1986;
http://www.ecology.com/2013/08/27/global-impact-
feral-cats/); therefore, the genetic diversity of the
domestic cat is likely to be very high (OʼBrian et al.,
1993; Lipinski et al., 2008). Here, male competition
during the breeding period, including sperm competi-
tion, is expected to be very intensive due to the lifestyle
of this species (social lifestyle, high population density
related to anthropogenic factors, and promiscuity)
(Izawa et al., 1982). Males with a low sperm quality
should have poorer opportunities for procreation in
this case.
We tested this hypothesis in a series of experiments
that addressed the effect of male competition and
sperm quality (percentage of morphologically intact
spermatozoa) on reproductive success in the domestic
cat. We assumed that the number of males mating with
a female would affect the fertility of the latter. More-
over, we expected that males with a higher sperm qual-
ity would have a reproductive advantage.
MATERIALS AND METHODS
The studies were performed at the Chernogolovka
experimental research station of IEE RAS in 2011–
2015. Domestic cats, with the exception of nursing

756
 EFFECT OF THE NUMBER OF MATING PARTNERS AND SPERM QUALITY
females, were housed individually in cages (2 × 1 × 2 m
in size) almost year round. Females and males were
housed together during the rutting period only. The
daily food ration consisted of approximately 200 g
chicken meat supplemented with vitamins and miner-
als, and water was provided ad libitum.
Twenty-six sexually mature animals (11 males and
15 females) were used in the study. Sperm quality was
assessed before mating in all males used in the study.
Electroejaculation was used for sperm sample collec-
tion. A rectal probe of 1.65 cm in diameter with two
copper electrodes and a variable-voltage electroejacu-
lator from P_T Electronics (Oregon, United States)
were used in the study; the output voltage ranged from
2 to 6 V (Jewgenow et al., 2006; Erofeeva et al., 2014).
Ejaculate smears were prepared for morphological
characterization. Microscopic observation was used to
assess the content of morphologically intact (normal)
spermatozoa. The configuration of at least 200 sper-
matozoa was analyzed in each smear (Glukhov and
Naidenko, 2013). The males were assigned to terato-
and normospermic male groups after the assessment
according to the content of abnormal spermatozoa.
Six of the 11 males used in the experiments had normal
sperm quality (more than 40% intact spermatozoa in
the ejaculate), and five were teratospermic (less than
40% intact spermatozoa in the ejaculate).
Two mating schemes were used during the rutting
period for the assessment of male competition on
reproductive success in the domestic cat.
The first scheme (“mating without competition”)
implied pair housing of the animals (one female and
one terato- or normospermic male). The number of
copulations was not limited. The animals were housed
together for two days, on average. Animal behavior
was monitored by outdoor video surveillance cameras.
The number and duration of copulations were the only
parameters recorded during video analysis. Twenty-
two pairs were formed, and 18 litters were produced.
The second mating scheme was used for the assess-
ment of male and female reproductive success in
domestic cats in the case of male competition and for
the identification of female preferences in mating
partner choice. One male and one female were housed
together for 4 h, and then the male was removed.
Another male was placed together with the female
after 10–12 h, and the animals remained together
during 4 h. Each female used in these experiments was
introduced to one teratospermic male and one normo-
spermic male, but the order of males was reversed in
every two consequent experiments. Sixteen experi-
ments were performed according to this scheme. This
series of experiments involved visual monitoring with
continuous data registration, and therefore all types of
pairwise social interactions of the animals could be
identified. Friendly and aggressive contacts and sexual
behavior were paid the most attention during behavior
analysis (see Erofeeva, 2010, for details). This analysis
BIOLOGY BULLETIN Vol. 45 No. 7 2018
757
revealed differences in the females’ reactions to terato-
and normospermic males. Ten litters were obtained
when this breeding scheme was used.
The animals were housed together when the
females were in estrus, as inferred from the female
behavior. Behavioral estrus is usually conspicuous in
the domestic cat, as the females frequently rub their
cheeks and ears on objects and show lordosis and high
vocalization frequency. Regardless of the domestic
cat’s ability to litter up to three times a year, all matings
were performed only once a year (in spring) in order to
eliminate the putative effect of the season on the litter
size and to standardize the time intervals between lit-
ters in the females (Prescott, 1973; Nutter et al., 2004).
The same males (six normospermic and five ter-
atospermic) were used in the first and second series of
experiments. Every male was mated to a different
female during each consecutive mating period. If a
female was mated with a normospermic male in the
first mating scheme (mating without competition), a
teratospermic male was introduced to the female
during the subsequent mating period. If the female was
first mated with a teratospermic male and then with a
normospermic male in the second mating scheme, a
reversed order of mating (first with a normospermic
male) was used during the subsequent mating period.
Female reproductive success was inferred from the
number of kittens in the litter determined within one
day after birth. Male reproductive success was inferred
from the number of kittens born from this male and all
females mated with it. Paternity tests based on micro-
satellite analysis with Fca 733, Fca 723, Fca 731, Fca
441, Fca 736, Fca 742, Fca 740, Fca 749, Sry, F 124, F 85,
and F 53 primers (Mennotti-Raymond et al., 1999)
were performed at the facility of molecular diagnostics
methods (Severtsov Institute of Ecology and Evolu-
tion, Russian Academy of Sciences). The number of
litters obtained in the entire study was 28 (97 kittens).
Microsoft Excel and Statistica 6.0 software were
used to perform the statistical analysis. Data are pre-
sented in the graphs as mean values and standard
errors of the mean (M ± SE). Nonparametric methods
of data analysis were used in the statistical processing.
The Mann–Whitney U-Test was used to compare the
female reproductive success for the different mating
schemes. The Wilcoxon matched pairs test was used as
the reproductive success of terato- and normospermic
males and estrous female behavior towards the males
were compared.
RESULTS
Female Reproductive Success
Eighteen litters (54 pups (3 ± 0.3 pups per female))
were obtained from females mated with a single male,
and ten litters (43 pups (4.3 ± 0.5 pups per female))
were obtained from females mated with two males.
Female reproductive success differed significantly
758 EROFEEVA et al.

4
Average litter size

0
One male Two males

Fig. 1. Litter size for the different mating schemes.

Normospermic males Teratospermic males

4.5
4.0
Average number of offspring

*
3.5
3.0
2.5
2.0
1.5
1.0
0.5
0
In females mated In females mated
to one male to two males

Fig. 2. Reproductive success of normo- and teratospermic males for the different mating schemes.

between the two different mating schemes. The aver-
age litter size was significantly higher in females mated
with two males than in females mated with one male
(Mann–Whitney test, Z = 1.97, n1 = 18, n2 = 10, p < 0.05)
(Fig. 1). The partner sperm quality did not affect litter
size in females mated to a single male (Fig. 2).
order of mating with the female. In contrast to mating
of a single female to a single male, reproductive suc-
cess of normo- and teratospermic males differed when
a female was mated with two males: it was significantly
higher for the normospermic males than for the ter-
atospermic ones (Wilcoxon’s test for paired samples:
n = 10, Z = 2.3, p < 0.05; Fig. 2).

Male Reproductive Success Here, a significant reproductive advantage of nor-

The effect of the mating scheme on male reproduc-
tive success was more complex than in the females.
The number of kittens sired by each male varied con-
siderably and depended on the sperm quality and the
mospermic males was only observed when a normo-
spermic male was the first of the two males mated with
one female. The majority of 23 kittens born after five
experiments of this type were sired by the first, normo-
spermic male, whereas the teratospermic males that

BIOLOGY BULLETIN Vol. 45 No. 7 2018

 EFFECT OF THE NUMBER OF MATING PARTNERS AND SPERM QUALITY 759

5.0
4.5
4.0

Average number of offspring

3.5
3.0
2.5
2.0
1.5
1.0
0.5
0
1st male 2nd male
(normospermic) (teratospermic)
Fig. 3. Dependence of reproductive success of males on the order of mating to the female (the normospermic male was the first
to mate with the female).

3.5

3.0
Average number of offspring

2.5

2.0

1.5

1.0

0.5

0
1st male 2nd male
(teratospermic) (normospermic)

Fig. 4. Dependence of reproductive success of males on the order of mating to the female (the teratospermic male was the first to
mate with the female).

were the second to mate hardly ever procreated (Fig. 3,
Wilcoxon’s paired test: n = 5, Z = 2.0, p < 0.05). How-
ever, if a teratospermic male was the first to mate with
a female, no differences in reproductive success of
normo- and teratospermic males were observed
(Fig. 4; five litters (20 kittens) obtained in five experi-
ments; Wilcoxon’s paired test: n = 5, p = 1).
Female Behavior towards the Males
We did not observe mating order–dependent dif-
ferences in female behavior towards males. The fre-
quency of aggressive and affiliative behavioral displays
towards the first and second male were similar, and
the number of copulations with those males was
approximately the same.
The differences in female behavior towards the
partner were associated with male sperm quality. The
females preferred normospermic males: affiliative
behavioral displays towards these males were signifi-
cantly more frequent (Fig. 5; Wilcoxon’s paired test:
n = 10, Z = 2.0, p < 0.05), and the same was true for
the number of copulations (Fig. 6; Wilcoxon’s paired
test: n = 10, Z = 1.9, p < 0.05). One should note that
males from both groups were equally persistent in their

BIOLOGY BULLETIN Vol. 45 No. 7 2018

760
50
45
40
Behavioral acts per h activity
35
30
25
20
15
10
5
0
Normospermic male cats
Fig. 5. Affiliative behavior of the females towards normo- and teratospermic males.
mating attempts. There were no significant differences
between the frequency of behavioral manifestations
directed towards subsequent copulation (mounting
attempt, copulation attempt, stalking of the female,
and others) in normo- and teratospermic males
(12.1 ± 3.1 and 18 ± 3.1 instances per 1 h of activity,
respectively; n = 5, ns).
Multiple paternity (presence of kittens sired by dif-
ferent males in one litter) was observed only in 20% lit-
ters (n = 10) from females mated with two males.
Here, manifestation of multiple paternity in the litter
did not depend on the mating order.
DISCUSSION
The sex conflict theory predicts that male repro-
ductive success depends on the number of sexual part-
ners, whereas female reproductive success depends on
partner quality, but not on the number (Bateman, 1948;
Trivers, 1972; compare Vasilieva and Tchabovsky, 2015).
However, multiple copulations (polyandry, i.e., mat-
ing with several males) are quite common for females
from different animal taxa, including mammals (Birk-
head and Møller, 1998; Zeh, D.W. and Zeh, J.A.,
2001; Stockley, 2003; Simmons, 2005). Multiple cop-
ulations of females are usually associated with
improved quality and survival of the offspring (Keil
and Sachser, 1998; Zeh, D.W. and Zeh, J.A., 2001;
Stockley, 2003; Simmons, 2005; Fisher et al., 2006;
Andersson and Simmons, 2006; Firman and Sim-
mons, 2008). Onlya few studies have demonstrated an
increase in litter size after the mating of a female with
several males (Madsen et al., 1992; Olsson and Shine,
1997; Hoogland, 1998; Birkhead, 2000; Jennions and
Petrie, 2000; Zorenko and Motmiller, 2005; Erofeeva
EROFEEVA et al.
Teratospermic male cats
and Naidenko, 2012). Other studies did not reveal an
effect of partner number on litter size (Keil and
Sachser, 1998; Wolff and Dunlap, 2002; Firman and
Simmons, 2008; Lane et al., 2008).
The mating of domestic cat females with two males
evoked a 30% increase in the average litter size in the
present study; that is, mating with several partners
conferred a substantial advantage to the females.
However, the reasons for this enhancement of repro-
ductive success remain to be elucidated. An increase in
the copulation frequency and number in females
mated with two males could be one of the reasons
(Wolff and Dunlap, 2002; Simmons, 2005; Firman
and Simmons, 2008). For instance, an increase in the
copulation intensity was reported to enhance repro-
ductive success, ovulation intensity, and the number of
implanted embryos in the social vole (Microtus socia-
lis) (Zorenko and Motmiller, 2005). Moreover, the
very presence of another conspecific male was shown
to enhance the male reproductive efforts and copula-
tion intensity in the meadow vole (Microtus pennsyl-
vanicus) (del Barco-Trillo and Ferkin, 2004). In addi-
tion to elevated copulation frequency, the number of
spermatozoa in the ejaculate increased when males
mated with females under these conditions. Similarly,
the mating scheme we used could have increased the
copulation intensity in the domestic cat. For instance,
a similar effect of repeated mating separated by a cer-
tain time interval was observed in our studies of
another feline species, the Eurasian lynx (Lynx lynx)
(Naidenko et al., 2007; Erofeeva, 2010). We demon-
strated that high copulation efficiency was only
observed for a few hours after an estrous female was
introduced to a male, but if a female was introduced to
BIOLOGY BULLETIN Vol. 45 No. 7 2018
 EFFECT OF THE NUMBER OF MATING PARTNERS AND SPERM QUALITY
14
12
Average number of copulations
10
0
Normospermic male cats
Fig. 6. Number of the females’ copulations with normo- and teratospermic males.
two males with a 10–12 h interval between matings,
the copulation frequency was high in both matings.
Therefore, regardless of a larger number of copulations
in the first mating scheme (since the mating time was
not limited), the copulation intensity was lower.
The reduction of embryonic loss is another putative
reason for fertility increase in a female mated with two
males (Madsen et al., 1992; Olsson and Shine, 1997;
Zorenko and Motmiller, 2005). Indeed, higher repro-
ductive success in the Eurasian lynx mated with differ-
ent partners was largely due to a decrease in embryonic
loss (Erofeeva and Naidenko, 2012). Unfortunately,
embryonic loss monitoring in the present study was
impossible. The only approach for noninvasive detec-
tion of embryonic loss currently available involves
ultrasound assessment of corpora lutea numbers and
comparison of these data to the number of pups in the
litter. However, the equipment currently used by the
authors does not allow for precise corpora lutea count-
ing in small animals, such as the domestic cat.
Unfavorable living conditions and/or low genetic
diversity can also be reasons for higher reproductive
success upon mating with two males. For instance,
Bateman (1948) showed that mating with several
males could enhance female reproductive success
under unfavorable conditions. This phenomenon was
supposedly related to transfer of a number of nutrients
to the female organism with male sperm and the
resulting increase in clutch size on nutrient-depleted
media in the fruit fly (Drosophila melanogaster). An
increase in female reproductive success upon mating
with two partners in reptiles was only reported in pop-
ulations with a low genetic diversity, where it was asso-
ciated with a decrease in embryonic loss (Madsen
BIOLOGY BULLETIN Vol. 45 No. 7
761
Teratospermic male cats
et al., 1992; Olsson and Shine, 1997). Reproductive
success upon mating with one male did not differ from
that upon mating with two males if the genetic diver-
sity of the population was high (Madsen et al., 1992;
Olsson and Shine, 1997). However, these factors
(unfavorable conditions or low genetic diversity) are
unlikely to have affected litter size changes in females
mated with one or two partners in the present study.
The cats received abundant food, even though the diet
was not varied (200 g chicken meat per cat per day,
vitamin supplements, and one day of fasting per week).
The average litter sizes (3 ± 0.31 and 4.3 ± 0.47 pups per
female for the different mating schemes) are in good
agreement with published data (Hall and Pierce, 1934;
Deag et al., 1987).
No differences in the female’s reaction to males
that would depend on mating order or familiarity with
the male were observed when any female was mated
with two males. A preference for the first (“familiar”)
male was observed in our earlier studies of another
feline species (the Eurasian lynx) (Naidenko and Ero-
feeva, 2004; Erofeeva, 2010): the females accepted the
male faster and copulated more often (Erofeeva and
Naidenko, 2012). These differences between two
feline species are probably due to the species-specific
features of space use. The Eurasian lynx is a solitary
animal, similarly to most other felines. Females only
encounter males during the rutting period. However,
the domestic cat is more social and forms large colo-
nies at sites with a high population density (Izawa
et al., 1982; Yamane, 1998; Say et al., 2002). All males
that live in close proximity to the female will be
“familiar” to her in this case, and the order of mating
with these males will be irrelevant.
2018
762 EROFEEVA et al.
Here, we observed that the females’ behavior
towards normo- and teratospermic males was differ-
ent. The females showed affiliative behavior towards
normospermic males more often, and the frequency of
copulation with these males was higher. This situation
could be expected, as females are usually assumed to
prefer the most competitive males in order to enhance
reproductive success through mating (Gerlinskaya,
2009). Indeed, the majority of offspring from each
domestic cat female was sired by normospermic
males.
Let us now discuss the reasons for the females to
mate with less competitive teratospermic males. One
of the explanations for multiple matings in mamma-
lian females is the female’s ability to choose “better
genes” (or “more appropriate” genes) that can confer
higher viability and sexual attractiveness to the off-
spring (Keil and Sachser, 1998; Birkhead, 2000; Jen-
nions and Petrie, 2000; Stockley, 2003; Firman and
Simmons, 2008). Teratospermic males can also be
carriers of such genes. Females can influence male
sperm competition, and the assessment of partner
quality and genetic compatibility can occur in the
female’s reproductive organs after mating (“cryptic
female choice”) (Birkhead, 2000). The second expla-
nation implies a reduced risk of infanticide after mat-
ing with the second male. Infanticide is a serious prob-
lem for the domestic cat, especially at high population
densities (Macdonald et al., 1987; Natoli, 1990; Feld-
man, 1993). Mating of the female with several partners
for infanticide prevention was reported for many other
mammalian species (Wolff and Macdonald, 2004;
Klemme and Ylönen, 2010; Huchard et al., 2012).
However, we assume that the mating of domestic
cat females with less competitive teratospermic males
is involuntary and stems from sexual coercion, a phe-
nomenon very common among animals (Clutton-
Brock and Parker, 1995). Domestic cat males are, on
average, 30–40% larger than the females, and there-
fore, the males play a decisive role in the relationships
in animal pairs during the rutting period (Erofeeva
et al., 2015). The males “court” the females very per-
sistently, and hiding from the male is the female’s only
opportunity to avoid undesired copulation. The
female is often coerced into mating with several males
due to the males’ persistence (Alcock et al., 1977;
Clutton-Brock and Parker, 1995; Thonhauser et al.,
2013). An attempt to avoid injuries related to overly
intensive courtship can be an important factor that
affects the acceptance of a second male by the female
in this case (Birkhead, 2000).
Female behavior has only a slight effect on male
reproductive success in this situation, whereas the
male’s quality and reproductive efforts acquire consid-
erable importance. Most kittens born during the pres-
ent study were sired by normospermic males that had
a higher sperm quality (inferred from the relative
abundance of intact sperm, %) than the teratospermic
males. The dependence of male reproductive success
on the relative abundance of intact spermatozoa was
also demonstrated for other animals (Naidenko et al.,
2007; Erofeeva, 2011). However, significant differ-
ences in reproductive success of normo- and terato-
spermic cat males were only observed when one
female was mated with two males and the normosper-
mic male cat was the first to mate. Male success did
not otherwise depend on sperm quality. Teratospermic
males are believed to use compensatory mechanisms
that increase reproductive success: for instance, a
larger testicle volume (Neubauer et al., 2004), a lower
apoptosis level (Pukazhenthi et al., 2006), and a larger
absolute number of spermatozoa in the ejaculate as a
consequence (Howard et al., 1990; Pukazhenthi et al.,
2000, 2006). However, neither a larger testicle volume
nor a larger number of spermatozoa were registered in
the teratospermic males used in our study. More suc-
cessful procreation of teratospermic males that were
the first to mate with the female can be related to
another distinctive feature of the males. In addition to
higher sperm quality, normospermic male cats may
stimulate the female more efficiently, and this is espe-
cially important in the case of induced ovulation.
Induced ovulation is the predominant ovulation type
in the domestic cat, and therefore the male’s ability to
stimulate the female can be of extreme importance for
successful procreation. The male that stimulates the
female more actively during copulation is supposed to
sire the majority of pups (Lariviere and Ferguson,
2003). However, if sperm from a teratospermic male is
already present in the female’s reproductive system,
successful stimulation of ovulation by the second (nor-
mospermic) male partner can increase the first male’s
siring chances. The normospermic males used in our
experiments sired almost all kittens when they were
the first to mate with the female and half of the kittens
when they were the second to mate.
We detected multiple paternity in only 20% litters
from females mated with two males. However, pro-
miscuous breeding in feral cat populations reportedly
resulted in multiple paternity in 68–83% of all litters
(Yamane, 1998; Say et al., 1999). The low relative
abundance of litters with multiple paternity in the
present study may be related to the small number of
males mated to one female (only two), whereas the
majority of multiple-paternity litters were detected in
urban populations with a very high animal density,
where up to nine males could mate with one female.
The abundance of litters with multiple paternity in the
countryside (at a low population density) was low (not
higher than 22%) (Say et al., 1999).
Thus, mating with two partners increased litter size
for each domestic cat female. The females preferred
BIOLOGY BULLETIN Vol. 45 No. 7 2018
 EFFECT OF THE NUMBER OF MATING PARTNERS AND SPERM QUALITY
normospermic (that is, more competitive and success-
ful) males, but the partner choice was limited, and
therefore the females were apparently forced to acco-
modate less competitive teratospermic males. The
male reproductive success was apparently less depen-
dent on female behavior and preferences and more
dependent on the sperm quality and order of mating
with the female in this situation.
ACKNOWLEDGMENTS
The authors are grateful to Ekaterina Pavlova,
Dmitrii Glukhov, and Alla Glukhova for help with
animal care and material collection.
This study was supported by the Russian Founda-
tion for Basic Research (project no. 15-04-07845 A).
Funding for animal maintenance was provided by the
State Task “Collection of Live Animals of Wild Pred-
atory and Ungulate Species at the Chernogolovka
Research Facility, IEEP RAS” (project no. 0109-
2016-0002).
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Translated by S. Semenova