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Applied Animal Behaviour Science 113 (2008) 215223

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Sucking and allosucking duration in farmed

red deer (Cervus elaphus)
Jorga Drabkova a,b, Jitka Bartosova a,*, Ludek Bartos a,
Radim Kotrba a, Jan Pluhacek a, Ludmila Svecova a,
Adam Dusek a, Tomas Kott a
a
b

Institute of Animal Science, Pratelstv 815, 104 01 Prague Uhrneves, Czech Republic
Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences,
Kamycka 129, 165 21 Prague, Czech Republic
Accepted 30 October 2007
Available online 20 February 2008

Abstract
Sucking duration in ungulates does not only mean milk transfer, but is also associated with maternal care
in general. It seems to be a reflection of offspring demand rather than solely milk transfer rate. Thus, the
objective of this study was to discriminate between sucking and allosucking (i.e. sucking non-maternal hind)
behaviour in red deer according to the sucking duration.
We hypothesized that: (1) calves should suck longer from their mothers than allosuck from non-maternal
hinds; (2) sucking duration of calves frequently nursed by a particular non-maternal hind should be longer
than that of calves occasionally allonursed; (3) sucking duration should be longer for bouts including one
calf than two or more calves sucking simultaneously; (4) male calves should suck and allosuck longer than
female calves; and (5) primiparous hinds should nurse and allonurse longer than multiparous hinds. We
observed sucking behaviour of 25 hinds and their 38 calves (from birth until the youngest calf reached one
month of age) in two seasons. We recorded 1730 sucking bouts, of which 11.62% in the first season and
4.37% in the second season were non-filial. The duration of filial sucking was significantly longer than nonfilial sucking. A large individual variance in the incidence of non-filial sucking in both the calves and hinds
was found. Therefore, the non-filial hindcalf pairs were categorized in two clusters according to the
frequency of nursing non-filial calves for one hind in relationship to all nursing events for this hind by a
cluster analysis (PROC CLUSTER, SAS). We used a general linear mixed model, GLMM (PROC MIXED,
SAS) to test the influence of hind relationship to the nursed calf (filial, frequently allosucking non-filial, or
occasionally allosucking non-filial pair). Sucking duration of occasionally allosucking non-filial calves was
only marginally different from that of filial calves. There was no difference between the two groups of

* Corresponding author. Tel.: +420 267 009 598; fax: +420 267 710 779.
E-mail address: bartosova.jitka@vuzv.cz (J. Bartosova).
0168-1591/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.applanim.2007.10.009

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J. Drabkova et al. / Applied Animal Behaviour Science 113 (2008) 215223

non-filial calves. Multiple sucking bouts were shorter than those with one calf. Male calves sucked longer
than female calves; however, the greatest difference was recorded between frequently allosucking non-filial
pairs of both sexes. Frequently allosucking non-filial males sucked the longest and differently from
occasionally allosucking non-filial males. Frequently allosucking non-filial females sucked the shortest and
differently from filial calves of both sexes. It is more likely that allosucking seems to be more important for
male rather than female calves.
Therefore, it is concluded that allosucking calves differ in their sucking behaviour and two types of
allosuckers (frequent and occasional) should be taken into account when analyzing allosuckling behaviour.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Sucking behaviour; Sucking duration; Allosucking; Red deer; Cervus elaphus

1. Introduction
Sucking duration was previously used to estimate milk transfer, such as in fallow deer, Dama
dama and white-tailed deer, Odocoileus virginianus (Gauthier and Barrette, 1985). However,
neither sucking duration nor its frequency may be a reliable indicator of milk intake in fallow deer
(Birgersson and Ekvall, 1994) and ungulates in general (Cameron, 1998). Sucking duration serves
for nutrition and is also associated with general maternal care, e.g., young that are in dangerous or
stressful situations search for their mothers and are calmed by sucking them (Lee, 1987; French,
1998; Roulin, 2002). Sucking duration seems, rather, to be a reflection of offspring vital demands
(including safety, social contact, etc.) than solely milk transfer rate (Mendl and Paul, 1989).
Allosuckling, i.e. nursing of a non-filial offspring, has been reported in more than 100
mammalian species (Packer et al., 1992). Its causes are not yet fully understood. As suggested in
our previous papers on red deer, Cervus elaphus (Bartos et al., 2001a) and cattle, Bos taurus
(Vchova and Bartos, 2005), we assume that allosuckling has evolved as an altruistic behaviour of
females through which allosucking offspring compensate for deficiencies from their dams
(Landete-Castillejos et al., 2000), rather than as a females misbehaviour due to mistaken
recognition of progeny (Tiplady, 1990; Cassinello, 1999).
Sucking duration changes over the course of lactation. As young grow older, short sucking
duration often occurs, for example in red deer (Bubenik, 1965), white-tailed deer and fallow deer
(Gauthier and Barrette, 1985), bighorn sheep, Ovis canadensis californiana (Shackleton and
Haywood, 1985), and cattle (Lidfors et al., 1994). Such short sucking may allow the young to
gain some milk but may occur especially to reinforce the motheryoung bond (Lent, 1974;
Prescott, 1981; Shackleton and Haywood, 1985; Festa-Bianchet, 1988). A decrease in sucking
duration and frequency may also correspond to the beginning of alternative feeding as shown in
mouflon, Ovis musimon (Reale and Bousses, 1995), Saharan arrui, Ammotragus lervia
(Cassinello, 2001), and red deer (Vasquez et al., 2004). Alternatively, some authors have
observed in cattle (Reinhardt and Reinhardt, 1981) and llama, Lama glama (Gauly et al., 2003),
relatively constant sucking duration during the whole lactation.
Sex-differential sucking duration may be associated with sex-biased maternal care, directed
especially to males in polygynous dimorphic ungulates, such as the red deer (Clutton-Brock et al.,
1982). It could be hypothesized that this may also be the case for allosuckling. Data suggests,
however, that the sex of the calf did not play a role in either allosucking frequency in red (Bartos
et al., 2001a) and fallow deer (Pelabon et al., 1998), or allosucking duration in fallow deer (Ekvall,
1998; Pelabon et al., 1998), red deer (Vasquez et al., 2004), and Saharan arrui (Cassinello, 1999).

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217

Suckling behaviour depends closely on the age and parity of the mother (Reale et al., 1999).
Young mothers have to invest in both the care of their progeny and their own growth (FestaBianchet, 1988; Birgersson and Ekvall, 1994). Thus, they should display less intense maternal
behaviour as well as a lack of maternal experience. However, they may also not control the
demands of their young, and hence overinvest in them (Trivers, 1974). Primiparous
(Birgersson and Ekvall, 1994) as well as young mothers (Ekvall, 1998; Reale et al., 1999) fallow
deer and mouflon have been observed to allow long suckling. On the contrary, young ewes nursed
for shorter periods but much more frequently, than the older ones in bighorn sheep (FestaBianchet, 1988) and mouflon (Reale and Bousses, 1995). Primiparous dams may also have
problems recognizing their own offspring, as suggested for water buffalo, Bubals bubalis
(Murphey et al., 1995), Saharan arrui (Cassinello, 1999), and muskox, Ovibos moschatus
(Tiplady, 1990).
Allosucking duration was shorter than maternal sucking duration in cattle (Waltl et al., 1995),
fallow deer (Ekvall, 1998), and red deer (Vasquez et al., 2004). It was suggested that nursing
females may prefer certain non-filial individuals compared to other in fallow (Pelabon et al.,
1998; Ekvall, 1998) and red deer (Bartos et al., 2001b). However, it is unknown how this selective
allosuckling affects sucking duration.
In the present study, we focus on the difference between sucking and allosucking duration in
red deer. We have tested five hypotheses concerning the effect of calfs sex and age, number of
calves sucking in one bout, and parity of the nursing hind on allosucking duration: (1) calves
should suck longer from their mothers than from non-maternal hinds; (2) sucking duration of
calves frequently allonursed by a particular non-maternal hind should be longer than those in
occasionally allonursed calves; (3) sucking duration should be longer for bouts including one calf
than two or more calves sucking simultaneously; (4) male calves should both suck and allosuck
longer than female calves; and (5) primiparous hinds should have longer nursing and allonursing
durations than multiparous hinds.
2. Methods
2.1. Animals and handling
The study was performed during two seasons (2003 and 2005) on a group of 28 farmed red deer hinds at
Podlesko, an experimental facility belonging to the Institute of Animal Science (Czech Republic). Fifteen
and 23 hinds gave birth to a calf in respective seasons, 6 and 4, respectively, being primiparous. Fourteen
hinds gave birth in both seasons.
The long-term socially stable group contained hinds born between 1992 and 2002 and have been living
together as one herd since 2000 (one of the dams was sold between observational seasons, no new animal
was introduced). Stags joined this group only during the rut (SeptemberNovember). The hinds and calves
were held in one of six enclosures with a 0.75 ha pasture. Animals grazed ad libitum during the growing
season (from April to October, covering lactation of the females) and were fed a supplementary grain
mixture and hay throughout the year.
In 2003, 10 male and five female calves were born between May 30 and June 16. In 2005, 13 male and 10
female calves were born between May 17 and June 27. No twins were born.
Hinds and calves were marked with coloured plastic collars. During the calving season, the newborns
were sought out every morning by caretakers walking through the enclosure. Each calf that had already dried
off was weighed with a hanging scale, marked with a collar, ear-tagged and its sex was determined.
Desertion by mothers resulting from interference at handling has never been observed in this study. For
verification of maternity DNA analysis was performed.
The study received approval for animal use and care from the institutional committee.

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2.2. Behavioural observation and definition

Observations started each season when the first calf was born and lasted until the youngest calf reached
one month of age. During each observation session, the animals were observed by one of 10 trained
observers. These observers watched with binoculars from a moveable, elevated observation tower, which
was placed by the fence of the enclosure. Sucking duration was measured using a stopwatch.
The data were collected using the ad libitum sampling method (Altmann, 1974). Each observation was
carried out to cover most of the sucking that occurred in the herd during the observation session. According
to our previous study (Illmann et al., 1997), the animals were observed during two sessions a day: from 5:00
to 8:00 a.m. and from 5:00 to 9:00 p.m. Ten observation sessions per week were performed.
For each observed sucking bout, we recorded the identity of the animals, the number of calves sucking the
hind at the time and the sucking duration. We considered a sucking bout to occur when one calf took a teat in
its mouth for at least 5 s and included all periods of sucking activity as long as the calf was not interrupted for
more than 1 min (Bartos et al., 2001a). One nursing bout was defined as nursing one or more calves by one
hind continuously (Waltl et al., 1995); a criterion of no interruptions of more than 1 min was met as well. When
another calf was sucking the hind simultaneously, the sucking bout was termed multiple.
2.3. Statistical analysis
All of the data were analyzed using the SAS System V 9.1 (SAS Inst. Inc., Cary, NC).
A cluster analysis was applied to categorize non-filial hindcalf pairs according to their allosucking
frequencies (the clustered variable was frequency of nursing non-filial calves for one hind in relationship to all
nursings for this hind, PROC CLUSTER). The non-filial hindcalf pairs from the cluster with a higher
proportion of allosuckling between the hind and non-filial calf from the whole number of nursings observed for
the hind were labelled as frequently allosucking (ranging from 0.07 to 0.28; 5 and 4 pairs in 2003 and 2005,
respectively); the second cluster contained occasionally allosucking hindcalf pairs for those in which
allosucking rarely occurred (ranging from 0.01 to 0.06; 32 and 32 pairs in 2003 and 2005, respectively).
As neither sucking nor allosucking duration were normally distributed, differences between them were
tested using the nonparametric KruskalWallis test for location and scale differences (PROC NPAR1WAY).
Then the dependent variable sucking duration was transformed by BoxCox transformations (Timm and
Mieczkowski, 1997) to normalize the data. The effect of the tested factors (listed below) on sucking duration
was then assessed by fitting a general linear mixed model (PROC MIXED, SAS). To account for the
repeated measures, the analysis was performed using the interaction hind*calf as a random factor. After
visual inspection of the data (plotting the sucking duration for each hind in both seasons), the hinds, which
were observed in both observing seasons (N = 14), were treated as independent subjects. The full model
contained the fixed factors relationship of the hind to the nursed calf (filial/frequently allosucking nonfilial/occasionally allosucking non-filial pair), sex of the calf (male/female), age of the calf during
nursing, number of calves sucking in one bout (one/more), parity of the nursing hind (primiparous yes/
no), season (2003 or 2005), and considerable first-order interactions of these main factors. We used leastsquares-means (called hereafter adjusted means) instead of the arithmetic mean due to unbalanced
design. These adjusted means are, in effect, within-group means appropriately adjusted for the other factors
in the model. For multiple comparisons we used the TukeyKramer adjustment.

3. Results
Over the two seasons, 1730 sucking bouts in 1696 nursing bouts were recorded in 38 calves
born to 23 hinds. Of these sucking bouts, 11.62% (season 2003) and 4.37% (season 2005) were
directed towards non-maternal hinds (N = 128). At least one allosucking event was recorded for
26 of 38 calves. Four hinds did not allonurse at all. Allosucking occurred already from the first
week of the calfs life.

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219

The duration of filial sucking (83.25  1.81 s, mean  SE) was significantly longer than nonfilial sucking (67.52  7.22 s; x2(1) = 21.02, p < 0.0001). However, the duration of both, filial
and non-filial sucking bouts, displayed similar ranges, from 5 to 540 s, and from 5 to 541 s,
respectively. We also found a large individual variance in the incidence of non-filial sucking in
both the calves and hinds. Therefore, we categorized non-filial sucking according to sucking
frequencies in non-filial hindcalf pairs.
We recorded 1644 nursing bouts when only one calf nursed, and 43, eight and one cases when
two, three or four calves, respectively, were sucking simultaneously. Because of the low number
of multiple sucking bouts including more than two calves, these bouts were grouped and treated
as a single category.
The final general linear mixed model fitted for sucking duration transformed by BoxCox
transformations contained the following factors: age of the calf (F (1,1713) = 85.04; p < 0.0001),
relationship between the calf and the nursing hind (F (2,1557) = 2.61; p = 0.07), sex of the calf
(F (1,1506) = 6.40; p < 0.05), number of calves (F (1,1649) = 7.18; p < 0.01), and the interaction
relationship*sex (F (2,1075) = 9.60; p < 0.0001). The results showed that sucking duration
decreased with the increasing age of the calf.
In detailed comparison (adjusted means), the difference between sucking duration of filial and
occasionally allosucking non-filial hindcalf pairs was strongly indicated ( p < 0.03). The
sucking duration of frequently allosucking non-filial pairs lasted longer than that of occasionally
allosucking non-filial ones. However, this difference was non-significant ( p = 0.3). There was no
significant difference between the sucking duration of filial and frequently allosucking non-filial
pairs (Fig. 1).
The duration of sucking bouts where there was only one calf sucking were longer lasting
(5.90  0.11 s, adjusted means  SE) than those sucking bouts with multiple calves
(5.37  0.20 s; p < 0.01).
The sucking duration of male calves was longer than that of female calves (Fig. 2). However,
the interaction relationship*sex showed that the difference between the sexes was only
significant for the frequently allosucking non-filial pairs (Fig. 3). The sucking duration of the
frequently allosucking non-filial female calves was shortest compared to all of other categories. It
was significantly shorter than the sucking duration of the frequently allosucking non-filial male

Fig. 1. Comparison of sucking duration (adjusted means  SE) among filial, frequently and occasionally allosucking
non-filial hindcalf pairs (data transformed using BoxCox transformation).

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Fig. 2. Comparison of sucking duration (adjusted means  SE) between male and female calves (data transformed using
BoxCox transformation).

Fig. 3. Comparison of sucking duration (adjusted means  SE) among filial, frequently and occasionally allosucking
non-filial calves according to their sex (data transformed using BoxCox transformation).

calves and filial calves of both sexes. On the other hand, the sucking duration of the frequently
allosucking non-filial male calves was the longest. It was significantly longer than the frequently
allosucking non-filial female calves and occasionally allosucking non-filial male calves.
The factors parity of the nursing hind or season, as well as interaction terms not
mentioned above in the results, were not significant.
4. Discussion

Hypothesis 1. Calves should suck longer from their mothers than from non-maternal hinds.
The results indicated that filial sucking bouts were longer than non-filial ones, which is

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221

similar to observations reported (Waltl et al., 1995; Ekvall, 1998; Vasquez et al., 2004).
However, the duration of both, filial and non-filial sucking, displayed similar ranges that
indicated large individual variability or probably covered different types of allosuckers. For
that reason, we focused also on the sucking frequency and found very obvious and large
individual variances in the incidence of non-filial sucking in both calves and hinds. Further,
non-filial sucking bouts did not seem to be a homogenous group. The cluster analysis showed
two categories of allosuckers according to their frequency of sucking a particular nonmaternal hind: frequently and occasionally allosucking calves. Thus, two types of allosuckers
should be taken into account when assessing differences in sucking and allosucking
behaviour.
Hypothesis 2. Sucking duration of calves frequently allonursed by a particular non-maternal
hind should be longer than those in occasionally allonursed calves. After categorizing the nonfilial hindcalf pairs, we found no significant differences in sucking duration between filial and
frequently allosucking non-filial pairs. Filial calves, however, tended to have longer sucking
bouts than occasionally allosucking non-filial calves. According to these findings, we could
hypothesize that, compared to occasional allosuckers, frequently allosucking non-filial calves
could have similar advantageous food offerings by nursing hinds as filial calves have from their
mothers. Thus, we can presume the frequently allosucking calves could be preferred by the hinds,
while occasionally allosucking calves may not be preferred, as was previously suggested for
fallow (Pelabon et al., 1998; Ekvall, 1998) and red deer (Bartos et al., 2001b). Further research is
needed in this area.
Hypothesis 3. Sucking duration should be longer for bouts including one calf than two or more
simultaneously sucking calves. It was found that the number of calves taking part in a sucking
bout affected the duration of sucking. For those bouts involving only one calf, the sucking
duration was longer than those consisting of multiple calves. This finding may reflect limited
maternal investment by the nursing hind. Lactation is a very demanding maternal activity
that can often decrease the condition or even the survival prospects of the nursing hind. All
but two of the multiple sucking bouts did not include the filial calf. Due to the low number
of multiple sucking bouts, we could not test the difference between the bouts including
frequent or occasionally allosucking non-filial calves. The duration of multiple sucking bouts
may also be influenced by the order in which the calves start sucking. We did not, however,
collect enough data to evaluate this factor, and therefore we encourage future research to
investigate this.
Hypothesis 4. Male calves should both suck and allosuck for longer than female calves. This
assumption was supported; however, in subsequent analysis a significant within-class difference
was only recorded between frequently allosucking non-filial male and female calves. These
results may indicate that allosucking acts as a compensation for inadequate nutritive supply, is of
higher importance for male than female calves. Differences in maternal investment in male and
female calves are probably caused by differences in male and female needs rather than an
adaptive trait of the hind (Clutton-Brock, 1991).
Hypothesis 5. Primiparous hinds should have longer nursing and allonursing durations than
multiparous hinds. More than one-third of the hinds in our study were primiparous. Still, in
contrast to our presumption based on research in other ungulates (Gauthier and Barrette, 1985;
Tiplady, 1990; Birgersson and Ekvall, 1994; Murphey et al., 1995; Reale et al., 1999; Cassinello,
1999), we did not find any significant effect of parity on either sucking or allosucking duration.

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5. Conclusion
Sucking duration differed between filial and non-filial calves. However, two types of
allosuckers (frequent and occasional) were detected according to their frequency of sucking a
particular non-maternal hind. Higher incidence of allosuckling within non-filial hindcalf pair
mostly led also to longer sucking than in those with rare allosuckling. Thus, two different tactics
of sucking non-maternal hinds seem to be involved. Therefore, using just filial and non-filial
classification of sucking alone most likely does not reflect causes and function of allosuckling.
Acknowledgements
We are grateful to Vratislav Ksada, Petr Janovsky and Anna Toszogyova for excellent field
assistance, Jose Panama for technical support, and Trevor DeVries and Ivo Keltner for improving
the English of the manuscript. The project was supported by the Ministry of Agriculture of the
Czech Republic (MZe 0002701402) and by the Czech Science Foundation (523/03/H076).
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