Behaviour 154 (2017) 115–129 brill.

com/beh

Giraffe nursing behaviour reflects

environmental conditions

Markéta Gloneková a , Pavla Jůnková Vymyslická b ,

Magdalena Žáčková a and Karolína Brandlová a,∗
a
Faculty of Tropical AgriSciences, Czech University of Life Sciences Prague,
Kamýcká 129, 165 21 Prague 6–Suchdol, Czech Republic
b
Faculty of Environmental Sciences, Czech University of Life Sciences Prague,
Kamýcká 129, 165 21 Prague, Czech Republic
* Corresponding author’s e-mail address: [email protected]

Accepted 3 November 2016; published online 16 November 2016

Abstract
Nursing behaviour is one component of mammalian life history strategy that demonstrates context-
dependent flexibility within species. Wild giraffes live in groups and feed themselves over large
area. In the zoo, giraffes are kept in enclosures with abundant food resources. We aimed to analyse
whether the nursing behaviour of giraffes differs between the zoo and nature reserve and discuss
which factor (food intake, presence of predators, population density) can explain the difference.
We observed seven and four female–calf pairs in the Bandia reserve, Senegal, and in Prague Zoo,
Czech Republic, respectively. Nursing bouts were less frequent and longer and calves were less
successful in nursing solicitations in the reserve, as females there seemed to be more selective for
nursing times and locations, likely due to presence of predators and differences in food intake.
Allonursing occurred more frequently in the zoo which can be attributed to higher population
density in captive conditions.

Keywords
Giraffa camelopardalis, nursing, suckling, allonursing, milk-theft, hider.

1. Introduction
Wild and captive behavioural comparisons provide opportunities to assess
the influence of proximate factors, helping to understand the behavioural
plasticity of species (Komers, 1997). This is especially interesting in the case
of nursing behaviour belonging among the most energetically costly parts
of parental investment (Clutton-Brock, 1991), and requiring well-balanced

© Koninklijke Brill NV, Leiden, 2017 DOI 10.1163/1568539X-00003413

116 Influence of environment on giraffe nursing

trade-offs in terms of resource partitioning. From an evolutionary point of

view, a species’ life history and specific adaptations influence the suck-
ling behaviour (Ralls et al., 1987). These ultimate factors, in response to
the availability of resources and presence of predators, formed the ‘hiding’
or ‘following’ rearing strategies in ungulates, which influence nursing fre-
quency and duration (Lent, 1974). ‘Hiding’ calves are only nursed a few
times a day with longer suckling bouts, as in roe deer, Capreolus capreolus,
nursing for 20–40 min (Rosenblatt et al., 1985), while ‘following’ calves
may suckle more often with shorter bouts, as in caribou, Rangifer tarandus,
nursing 50 s only, 30 times per day (Parker et al., 1990).
The general nursing pattern is influenced by proximate factors, which
comprise current social and environmental conditions. These conditions dif-
fer considerably between wild and captive populations, and include food
intake, presence of predators and actual population density (Réale et al.,
1999; Therrien et al., 2008; Pluháček et al., 2010).
Animals face decisions on how to allocate all activities to ensure their
survival and fitness (Illius et al., 2002). Optimal time allocation for foraging
behaviour may be constrained by many factors, forcing animals to adopt
compromises to meet their daily energetic needs (Hamel & Côté, 2008).
Wild ungulates, including giraffes, spend quite a large part of the day feeding
themselves (Pellew, 1984a, b). Adult giraffes spend approximately half of the
24 h per day feeding (Pellew, 1984a).
To compensate for high energy expenditures, lactating females are ex-
pected to modify their foraging behaviour to increase nutrient intake (Han-
well & Peaker, 1977; Bunnell & Gillingham, 1985; Ruckstuhl & Festa-
Bianchet, 1998), which can be achieved by foraging even longer (Shipley
et al., 1994). However, lactating females face a fundamental trade-off be-
tween foraging, vigilance (Illius & Fitzgibbon, 1994; Cowlishaw et al., 2004)
and nursing (Rachlow & Bowyer, 1998; Toïgo, 1999; Hamel & Côté, 2008;
Pluháček et al., 2010). High population density may help individuals to
decrease time spent on vigilance. However, this could also be a factor ex-
plaining existence of allonursing, nursing of non-filial offspring (Packer et
al., 1992). Under high population densities, so called ‘milk-theft’ is among
the most represented hypotheses explaining allonursing. In giraffes, this is
usually connected with suckling in a parallel position which prevents the
calf’s identification by the nursing female (Brandlová et al., 2013; Gloneková
et al., 2016).
 M. Gloneková et al. / Behaviour 154 (2017) 115–129 117

Giraffe calves have been described both as ‘followers’ (Kitchen, 1977;

Horwich et al., 1983) and ‘hiders’ (Langman, 1977). Ralls et al. (1986)
ranked the giraffe in an ‘intermediate’ cluster, suggesting their ability to
react to changing environmental conditions. The nursing duration in gi-
raffes is highly variable (4–360 s; Langman, 1977; Pratt & Anderson, 1979;
Nakamichi et al., 2015) and the reason for this is unknown. Giraffe calves are
born year round (Bercovitch & Berry, 2010). Females typically give birth to
one calf and rarely have twins (Foster & Dagg, 1972). Wild females leave the
herd before parturition and usually give birth alone (Estes, 1991). A week or
two after parturition, the calf mostly hides and the mother feeds apart, with
nursing occurring several times a day (Langman, 1977). After the hiding
period, the mother and calf join other females with offspring (Pratt & An-
derson, 1979). Calves of similar age form nurseries, guarded by one female,
while the other mothers are feeding (Pratt & Anderson, 1979). Giraffe moth-
ers in Bandia reserve spent more than 60% of time accompanied by the filial
calf (Malyjurková et al., 2014).
Captive giraffes are also typically separated before parturition and give
birth alone (EAZA Giraffe EEPs, 2006). The time when they return to the
herd depends on husbandry management decisions. Nurseries are less ob-
vious as the space is limited, food provided, and females are constantly
associated with the calf (Greene et al., 2006). In captivity, the time spent
feeding is shorter (Veasey et al., 1996), and the feeding pattern is markedly
changed compared to wild giraffe (Baxter & Plowman, 2001). Allonurs-
ing was recorded in giraffes more frequently in captivity (Gloneková et al.,
2016) than in the wild (Pratt & Anderson, 1979). This pattern of allonurs-
ing occurrence corresponds with other ‘hiders’, such as the common eland
Taurotragus oryx (Hejcmanová et al., 2011) and species with an ‘intermedi-
ate’ strategy such as the camel Camelus bactrianus (Brandlová et al., 2013).
Explanations for allonursing in giraffes are variable, oscillating between re-
ciprocal help among females sharing nursing duties in nurseries and the
milk-theft theory (Gloneková et al., 2016).
We aimed to find out whether the nursing behaviour of giraffes differs
between the nature reserve and the zoo in an attempt to assess the plasticity of
giraffe nursing behaviour. The next aim was to discuss possible explanations
of such differences with specific regard to food intake, presence of predators
and population density.
118 Influence of environment on giraffe nursing

2. Materials and methods

2.1. Study area
In Senegal, introduced Cape giraffes (Giraffa camelopardalis giraffa) were
observed in the Bandia reserve, a fenced area run for the purpose of safari-
tourism which contributes to the protection of the environment and wildlife
conservation at the same time. Encompassing an area of approximately
3500 ha, the reserve was created on the southwest boundary of the ‘Classi-
fied Forest Bandia’ by the Society for Protection of Environment and Fauna
in Senegal (SPEFS) in 1990 (Nežerková et al., 2004). Based on phytogeog-
raphy, the reserve belongs to the Sudan-Sahelian area (Antonínová et al.,
2004). The landscape is formed by plains with baobabs and dense shrubs
that make animal observation difficult. The vegetation is composed of a va-
riety of acacia species (Hejcmanová et al., 2010), primarily Acacia seyal
(Antonínová et al., 2004), the main nutrition source for giraffes. Two sea-
sons can be distinguished: the dry season (from November to June) and the
rainy season (from July to October; White, 1983; Lawesson, 1995). Fauna in
the Bandia reserve includes indigenous Senegalese species, such as African
buffalo (Syncerus caffer brachyceros), roan antelope (Hippotragus equinus
koba) and Western Derby eland (Taurotragus derbianus derbianus), as well
as animals introduced from South Africa, including common eland (Tauro-
tragus oryx oryx), impala (Aepyceros melampus), greater kudu (Tragelaphus
strepsiceros), and giraffes (Giraffa camelopardalis giraffa; Al-Ogoumrabe,
2002). Predators are represented by golden jackal (Canis aureus) and spotted
hyenas (Crocuta crocuta).
A captive herd of Rothschild’s giraffes (Giraffa camelopardalis roth-
schildi) was observed in Prague Zoo, where all giraffes were kept in a tem-
perate stable during most of the winter months. The giraffes spent most warm
days in an outdoor enclosure (2 ha), sharing the space with other species
including common eland (Taurotragus oryx spp.), addax (Adax nasomac-
ulatus), beisa oryx (Oryx beisa), red lechwe (Kobus leche cafuensis) and
Grevy’s zebra (Equus grevyi). Giraffes were fed ad libitum with alfalfa hay
and with limited amounts of branches and supplements of granulated food,
fruit and vegetables.
2.2. Studied animals
In the Bandia reserve, seven females with one calf each from a total of 28
giraffes present in the reserve, whose ancestors were introduced from South
 M. Gloneková et al. / Behaviour 154 (2017) 115–129 119

Africa in 1997, were observed during the dry season. The spacious area of
the reserve enables giraffes to behave naturally, thus breeding occurs non-
seasonally, females leave the herd for parturition and births occur without
human intervention. Following the reserve’s requirements, all observations
were conducted from a vehicle. During the first week of observations, each
individual was identified according to the unique skin pattern, body size,
shape of the horns, and sex (Estes, 1991). Photographs of each giraffe were
taken from both sides, and ID cards were created to facilitate the identifica-
tion. Mother–calf dyads were identified by maternal behaviour, particularly
nursing.
In Prague Zoo, we observed four females with one calf each (from a total
of 13 giraffes present in the herd). Females at advanced stages of pregnancy,
during parturition and shortly after parturition were separated and fed with
milk supplements. All calves in the herd were sired by the same bull, making
them half-siblings to one another. Giraffes were identified individually as
above.
For our study, giraffes were considered calves up to the age of about 18
months, when they generally stop suckling and leave their mother (Pellew,
1984a). Calves of various ages were present in both types of environments.
While in the zoo exact data of calf ages were known, this was not the case
of reserve animals, where we only divided them into smaller ones and taller
ones, for better orientation, according to their height or other signs as residue
of the umbilical cord (Table 1). We suppose that the smaller calves were 0–6
months old while the elder ones where labelled as taller.
2.3. Data collection
In Senegal, the data were recorded between 10 January and 13 March 2013.
Due to climatic conditions, data collection was generally conducted during
two sampling periods each day. The morning period started at 08:00 h, with a
break at noon starting at 12:00 h corresponding to the temperature peak, and
the afternoon sampling period started at 15:00 h and continued until dusk.
In the zoo, the observations were conducted every 14 days between January
and October 2011. Each sampling period lasted for 6 h. During the sampling
period, all visible suckling solicitations were recorded ad libitum.
A suckling solicitation included either suckling attempt or successful
nursing bout. A suckling attempt was defined as the act when the calf took
the teat into the mouth for less than 5 s (Drábková et al., 2008), or when
120 Influence of environment on giraffe nursing

Table 1.
The overview of observed calves in the two environments.

Name Environment Sex Age (months) Age category

Doubravka Zoo F 1–9

Frantisek Zoo M 3–12
Kuba Zoo M 5–12
Vilma Zoo F 0–9
Bertik Nature reserve M Smaller
Dusan Nature reserve M Taller
Lenka Nature reserve F Smaller
Majda Nature reserve F Smaller
Marketa Nature reserve F Taller
Ruzenka Nature reserve F Smaller
Vilem Nature reserve M Taller

the calf just approached the udder of the female closer than the length of
the head of the calf (Špinka & Illmann, 1992). For the definition of nursing
bout, we followed Drábková et al. (2008), who described it as each situation
when the calf held the teat in the mouth for 5 s or longer, irrespective of the
number of calves simultaneously participating in one nursing. The nursing
bout was considered finished when the calf stopped suckling for more than
10 s (Drábková et al., 2008).
For each solicitation, we recorded identity of the nursing female, iden-
tity of the suckling calf, sex of the calf, age category of the calf (smaller,
taller), position (antiparallel, parallel, perpendicular; Bartoš et al., 2001; Za-
pata et al., 2009; Figure 1), initiation (female/calf), relationship between the
suckling calf and nursing female (filial/non-filial), nursing duration (in sec-
onds), activity of the nursing female (feeding/watching surroundings/other),
and environment (zoo/reserve). It should be noted that the antiparallel and
perpendicular positions are thought to enable the lactating female to identify
the calf (Packer et al., 1992; Bartoš et al., 2001; Brandlová et al., 2013), while
the parallel position does not allow the female to recognise the calf (Bartoš
et al., 2001; Zapata et al., 2009; Brandlová et al., 2013). We recorded the ac-
tivity of the female just before and during nursing. We concluded that giraffe
is vigilant while the recorded activity was ‘watching surrounding’ (Cameron
& du Toit, 2005), in contrast to the recorded activity ‘feeding’ and ‘other’
(meaning walking, drinking, defecating, urinating, licking the calf), when
we considered the female non-vigilant.
 M. Gloneková et al. / Behaviour 154 (2017) 115–129 121

Figure 1. The schema of nursing positions (1, antiparallel position; 2, perpendicular position;
3, parallel position). Direction of the arrow indicates the head of animal.

2.4. Data analyses

Data were analysed with the STATISTICA 12 software package (StatSoft,
Tulsa, OK, USA). First we calculated the suckling success for each female-
calf pair (number of nursing bouts/total number of solicitations). As the data
were not normally distributed we used generalized mixed models (GLZ) to
analyse differences in suckling duration and in suckling success. For suck-
ling duration, we tested the influence of the independent categorical vari-
ables: position, sex of the calf, age category of the calf, relationship between
the suckling calf and nursing female, initiation, activity of the nursing fe-
male and the environment. As the analysis of factors influencing suckling
success did not allow us to use case specific variables, we used just those
which were not changing among different cases. Therefore, we reduced the
tested factors to: sex of the calf, relationship between the suckling calf and
nursing female, activity of the nursing female and the environment. We used
contingency tables to show the differences in suckling position in the two
environments.

3. Results
In the nature reserve, we recorded in total 141 solicitations during 22 days;
10 of the solicitations included a non-filial calf (7.1%). Solicitations resulted
in 13 nursing bouts (9.2%), and none of the nursing bouts were non-filial.
The mean nursing bout duration ± SE was 47.3 ± 12.6 s (range 25–67 s).
122 Influence of environment on giraffe nursing

Figure 2. The difference in nursing duration between two different environments.

In the zoo, we recorded 574 solicitations during 22 days; 118 of the so-
licitations included a non-filial calf (20.6%). Solicitations resulted in 120
nursing bouts (20.9%), of which 28 included non-filial calf (23.7% of all
nursing bouts). The mean nursing bout duration was 20.25 ± 17 s (range
5–67 s). The mean nursing duration was significantly longer in the reserve
(50.5 ± 3.69 s) than in the zoo (20.25 ± 1.57 s, W = 30.242, p < 0.001;
Figure 2). The nursing bout duration did not differ according to the sex of
the calf (W = 1.034, p = 0.31) nor according to the age category of the calf
(W = 0.063, p = 0.80) or other factors.
Regarding the number of suckling solicitations needed for a nursing bout,
the calves were more successful in the zoo (15%) than in the reserve (5%,
W = 4.49, p = 0.03; Figure 3). Calves in both environments tended to be
more successful when females were not feeding themselves. The nursing
bout was also longer in both environments when females were not feeding
themselves (W = 12.499, p < 0.01), however there was only one episode in
the reserve where the female was feeding herself while nursing.
Nursing bouts were more likely to occur in antiparallel (zoo) and perpen-
dicular (reserve) positions. Calves in the reserve never suckled successfully
in the parallel position (Figure 4).
 M. Gloneková et al. / Behaviour 154 (2017) 115–129 123

Figure 3. The difference in suckling success proportion of the calves between two different
environments.

Figure 4. The difference in suckling position of the calf between the two environments.
124 Influence of environment on giraffe nursing

4. Discussion
This study investigated the nursing behaviour, especially the differences, of
giraffes in two environments, a zoological garden and a fenced nature re-
serve. The data for this study were collected by the same observer using the
same methodology in both environments what makes the study unique. We
recorded significantly fewer suckling bouts in the reserve than in the zoo
during the same amount of time, as giraffes in the reserve are not forced to
stay within view of the observer and recording is limited by dense vegeta-
tion where even the tall giraffe may easily remain undetected. Despite this,
we revealed significant differences between general nursing patterns in the
reserve and in the zoo. We further discussed these differences with respect
to the food intake, presence of predators and population density in respective
environments and confirmed a behavioural plasticity of giraffes (Komers,
1997; Horová et al., 2015).
Calves in the reserve needed more solicitations for one successful nursing
bout than calves in the zoo. The lower suckling success rate in the reserve
was likely connected with a higher need for the mother’s vigilance in natu-
ral conditions, as the mother’s vigilance may directly contribute to offspring
survival (Lima & Dill, 1990; Hunter & Skinner, 1998). Females in the re-
serve seemed to reject the suckling solicitations of calves until they felt
safe, observed their surroundings and use the window of opportunity with
no predators or other disturbing factors. This may include also the presence
of the observer, despite the fact that the giraffes in the reserve are habituated
to the presence of cars.
The difference in nursing duration was striking, as nursing duration in the
reserve was almost twice as long as in the zoo. Longer nursing in the reserve
may be connected with lower nursing frequency and explained by differ-
ences in food intake, presence of predators, and population density. Mean
mother–calf distance in the reserve is larger than in the zoo, as in the reserve
the female searched for food and often left the calf alone (Malyjurkova et
al., 2014), suggesting that calves in the reserve have fewer opportunities for
nursing solicitations than calves in the zoo, which may result in an overall
lower nursing frequency. Moreover, we can expect that females in the re-
serve choose the right time and location to nurse their calves to minimise
the risk of predation. Consequently, when they find a proper time and place
they may try to take advantage of it which results in longer nursing dura-
tions. Higher population density in the zoo may also cause higher level of
 M. Gloneková et al. / Behaviour 154 (2017) 115–129 125

disturbance by other calves, which may make the suckling duration shorter.
Our finding corresponds with other studies describing longer nursing dura-
tions in the wild. Langman (1977) measured the nursing duration of giraffes
to be 45–360 s, and Pratt & Anderson (1979) measured the mean nursing
duration as 66 s, with 57 out of 415 nursing bouts taking more than 120 s.
In captivity, Nakamichi et al. (2015) observed giraffe nursing duration to be
approximately 60 s, with calves older than one month. Significantly longer
nursing bouts in the wild compared to captivity were also reported in other
ungulates such as camels (Brandlová et al., 2013) or goats (Bungo et al.,
1998).
In the reserve, the trade-offs between food intake and vigilance apparently
resulted in different nursing patterns than in the zoo. Despite a longer time
spent feeding in the reserve, the giraffes almost never fed themselves during
nursing, while feeding during nursing was relatively common in the zoo.
This was likely due to higher vigilance in the reserve where the giraffes
mostly carefully watched their surrounding during nursing, apparently to
search for predators or other danger. We recorded only one case when a calf
succeeded in suckling while the female was feeding herself in the reserve,
while in the zoo the calves often successfully suckled even if the females
were feeding themselves.
We also recorded differences in the suckling position of the calf. Calves in
the reserve suckled mostly in the perpendicular position and less suckling oc-
curred in the antiparallel position, while the parallel position was never used
for suckling. Calves in the zoo suckled most often in the antiparallel position,
but often also in the perpendicular and parallel positions. The parallel posi-
tion makes the identification of the calf for the female harder (Packer et al.,
1992). Nursing in the parallel position was also shorter. These results sug-
gest that females in the reserve were more focused on identification of calves
during nursing which may prevent non-filial calves from allonursing if ex-
plained by milk-theft (Gloneková et al., 2016). We assign those differences
to different population densities in respective environments. Higher popula-
tion density in the zoo can probably explain also the fact that in this study,
23.7% of nursing in the zoo was allonursing, while in the reserve we only
found allonursing attempts. Allonursing in wild giraffes has been inciden-
tally documented (Pratt & Anderson, 1979), while Gloneková et al. (2016)
recorded the highest allonursing rate among non-domesticated mammals in
captive giraffes with more than 80% of females allonursing and described
126 Influence of environment on giraffe nursing

in detail several hypotheses explaining allonursing. In our study, calves in

both environments attempted to suckle and to allosuckle in the parallel po-
sition, but only in the zoo did calves in this position succeed and suckle.
Such behaviour could be supported by lower vigilance of zoo females, which
were less concentrated on identification of suckling calves and also by lower
nutrition stress in captive females fed ad libitum. Long inter-individual dis-
tances and unlimited space enable lactating females to select a remote place
for nursing, which in combination with careful identification of the suckling
calf decreases opportunities for milk-theft. Allonursing in wild giraffes (as
documented by Pratt & Anderson (1979)) could therefore be explained by
an adaptive hypothesis.

Acknowledgements
We sincerely thank the Society for the Protection of the Environment and
Fauna in Senegal (SPEFS), Directorate of National Parks of Senegal (DPN)
and the management of the Prague Zoo for their support of the study. We
would also like to thank the NGO Derbianus Conservation, which provided
technical and logistical support during the research. We are also grateful
to Eva Baranyiová, Jan Pluháček, Luděk Bartoš and Radim Šumbera for
very useful comments. The study was supported by grants from the Czech
University of Life Sciences Prague, CIGA 20135010, CIGA 20134217 and
IGA 20165017.

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