Antipredator aspects of fallow deer bebaviour
during calving season at Doñana National Park (Spain)
C. SAN JOSÉ and F. BRAZA
Estación Bioldgica de Dotiana, CSIC Apdo 1056, 41080 Sevilla, Spain
Received 26 June
1990, accepted 1 March 1991
This study evaluates some morphological, ecological and behavioural aspects of
fallow deer mothers and fawns during the calving season in order to know the
antipredator tactics selected in this species living in a particularly open hahitat at
Doñana National Park. In this area (allow deer give birth in synchrony during the first
fortnight of June. At this time they move away from their matriarchal groups. A
concentration of births in the early afternoon was detected, coinciding with the
minimal activity of predators in the area. Mothers groom their offspring and ingest the
birth remains to prevent the attraction of predators. During the first days of life fawns
remain hidden keeping motionless.
KEY WORDS:
Dama dama, antipredator strategy, carving time, maternal behaviour.
Introduction
Materials and methods
139
140
Resulrs
143
Discussion
145
146
147
Acknowledgements
References
iNTRODUCTION
The fawns in many species of ungulates are specially liable to suffer from
predation during their first days of life. However, only a few works exist about the
mechanisms of developing an antipredator strategy at calving time, probably due to
the difficulty to observing mothers and their young at birth and during the days
immediately after birth.
Mothers and newborn young in those species use a ehiden> strategy (elk,
ALTMANN 1932; fallow deer, GILBERT 1968; Coke’s bartebeest, GosLiNG 1969;
bontebok, DAvID 1973, 1975; pronghorn, KITCHEN 1974; red deer, GIJINNESS et al.
1978; red buck, IRBY 1979). The species of these groups are basically classified as
*cbidero or ((followers> depending on whether the newborn lie concealed for their first
few clays or actively follow their mothers (LENT 1974, RAI.i.s et al. 1986). While
<<following>> has been viewed as a strategy for avoiding predators in open habitats,
hidinga is thought to reduce predation risk in closed habitats (LENT 1974, ESTES &
ESTES 1979).
As to the behaviour of fallow deer during the calving season, on the basis of the
studies we have conducted to date (ALvA.aEz et a!. 1975, BitAzA et al. 1988, SAN Jose
1988, BItizA & SAN Jose 1989), we may classify fallow deer in principle as ehider>’
since the young remain hidden the first days after birth and are only visited
periodically by their mothers, mainly to feed. Nevertheless, morphological, ecological
and behavioural aspects, which decide this assignment, have not been evaluated
quantitatively up to now. The aim of our study was to evaluate some of these aspects
in mothers and fawns during the calving season and to discuss the results in relation to
the antipredator strategies developed by this species and the physical characteristics
of the habitat.
MATERiALS AND METHODS
Observations included in this paper were carried out at Doñana National Park from 1982 to
1987. In this National Park of 73,000 ha (Fig. I) situated at the mouth of the river Guadatquivir,
the wild fallow deer population occupies an ecotone zone which corresponds to the transition
between the xerophyric schrub and the marshland (BRAZA 1975). In this ecorone we distinguish
between the following biotopes (according to ALLIER er al. 1974, and AGuIr.AR AMAT et al. 1979)
(Fig.1):
a) marshes: characterized by a strong seasonal dependence and made up of species like
Salicornia ramosissima, Arthrocnemum sp. and Scispus maritimus;
bI rush areas: form a mosaic wits pastures at the marshland borders and present a dominant
vegetation of macus marit&nus;
c) wet meadows: areas of pastures which are occasionally flooded and basically characterized
by the association of Amzena gaditana and 44sphodelus aestivus;
d) dry meadows: have a lower level of the subterranean water and present a characteristic
association of the species Urinea maritima and Anthemis cotula;
e) bracken areas: areas with a dense plant cover where the species Pseridium aquilinum
dominates;
f) schrubland: is made up of vegetation which colonized stabilized sands and is basically
characterized by species like Halirnium halimifolium, Cistus libanotis, Erica scoparia, Calluna vulgaris
and Stauracanthus genistoides, among which some isolated cork oaks (Quercus suber) are found.
At present, the Iberian lynx (Lynx pardina) is the only predator able to kill half-grown deer in
Doñana and, as we know, the lynx has very little influence on the early mortaliry of fallow deer
fawns (BELTRAN et al. 1985). Of other potential predators of f allow deer calves (fox, genet,
ichneumon, wildboar, Imperial eagle) rhe analyses do not reveal significant deer presence in their
diet (VENERO 1982, PAL0MARE5 1986, RAu 1988, FERRER 1989).
During 4 years (1983-1986), from the 25th of May to the 20th of June we conducted a census
by car at least once a week a fraction of the ecotone area which measured 5.5 km long and 1.5 km
wide (Fig.1).
Generally two observers record the number of solitary females (animals char were ar least 50
away from any other fallow deer). The proportion of solitary females was calculated with respect ro
the rotal number of females present in the area the same day.
The mean population size during the period 1983-1986 was 264.44 (* 7.89) deer wirh a mean
total number of females of 123.92 (±6.45). The mean calf-ratio was 60.37 (±2.05) (from BRAZA et
al. 1990).
The rutting season takes place during the first fortnight of October (BRAZA er al. 1986).
intipredator behaviour in fallow deer
ri
141
Area of Census
Fig. 1. Doñana National Park and study area (above). The main biotopes considered in the study arc
roportion ol the ecorone zone (below).
—
C. San José and F. Braza
142
198a
>
Ct
1984
0
U)
0
1985
30
-
20
10
L
I
25
May
Fig. 2
5
15
June
P
1986
I
25
5
July
Variations in the percentage of solitary females
Females give birth to a single fawn at the end of spring (BRAZA er al. 1988), after a gestation period
o 236 days (SAN Jos 1988),
During the calving season, in the area of census we searched daily to localize newborn fawns.
This was done on foot and sometimes counting on the assistance of a vehicle or a guard on
Antipredator behaviour in fallow deer
143
horseback. When we found a fawn we recorded all the environmental characteristics of the place
where it was hidden as well as the behaviour shown by the mother (if nearby) and the fawn at our
approach. Once captured the fawn was marked and measured (BRAZA et aI. 1988, SAN josI et al.
1989).
When it was not possible to detect the moment of birth we used various criteria to estimate
the condition vnewbornn, e.g. the presence of remains of the placenta, the still moist fur of the
fawn, the umbilical cord’s degree of healing, the hardening of the hooves and the presence of
cartilage on their points. These criteria have frequently been used to determine the age of newborn
fawns of other deer species (i.e. HAUGEN & SnAKE 1958).
Finally, in order to make more detailed observations, we followed in a ‘main observation area’
(aprox. 150 ha) the movements of the pregnant females. This control was carried out by means of a
binocular telescope (20(40) x (80(500) from a 30 m high observation tower continually from dawn to
nightfall.
We used non parametric statistics (SIEGEL 1956) to evaluate the level of significance of the
results.
RESULTS
A sudden increase in the proportion of solitary females was observed each year at
the beginning of June (Fig. 2). The distributions of solitary females in different years
are coincident (83 X 84, 84 X 85, 85 X 86, t = 1; P< 0.01; KendalL Coefficience of
Concordance). Only 108 birth dates of the 190 fawns captured could be known with
precision. The peak of solitary females coincides with the average date of birth
estimated. With the exception of 1982, with a mean date of birth the 5th of June, for
the rest of the years (1983-1987) most of births occurred within the 3 first days of
June: 1=1 (±3.7), isiS (±3.8), *=3 (±2.2), Yc=2 (±3.8) and i=2.5 (±4.3)
respectively.
In 41 cases it was possible to record exactly the time of birth. From these cases
we could calculate the average hour which is 14:16 (± 3.16). Most births (x = 25.54;
P’cO.OOi) were detected between 15:00 hr and 17:00 hr (Universal time) (Fig.3).
In all the births observed in their entirity (n = 20), the mother turns towards the
fawn, which Ties on the ground immediately after birth, and starts to lick it actively,
removing and ingesting the amniotic membranes which still cover the young. Finally,
D
0
C?
cy
CC
C
C)
C.)
a)
3
—
Distribution of births over the day’ight hourt (Universal time)
C. San José and F. Braza
144
rush
marsh
dry meadow
‘wet meadow
Ei
schrub
bracken
Habitat preference shown by femaLes for
Fig. 4.
hiding their fawn (n = 190).
the female ingests the newborn’s faeces. The newborn fawn has a dark brown coat
with white speckles on its back, the belly being entirely white. Its colour gets
progressively lighter in the course of the 1st week of life.
During the first days of life the fawn lies hidden. Its mother lies down some
metres off or goes away to join a group or to graze alone. The biotope of the hidding
place was not selected randomly (xi = 511.45; P<0.001). Most of the fawns
captured were found in a rush area Cx? = 467.46; P<O.0O1) (Fig.4). The fawn
remains motionless and adopts the typical flat-on-the-ground posture, frequently
curled up totally concealing its extremities and head, although the tendency to run
away when a person is approaching rapidly is correlated with the fawn’s age (r I;
P<0.001) (Fig. 5), increasing and surpassing a probability of 50% from the 3rd day
of life.
01190 fawns captured, for 89 it was possible to observe the mother’s behaviour
when we approached the fawn. In 76% of all cases the female went away from the
place where the young was hidden. In 28 cases the female stayed near the place where
the young was being marked at an average distance of 72.5 m (±45.2).
/
no flight
Itighi
lawn
Eig. 5.
age
(days)
TemporaL variation in the fawn’s reaction at the approach of a person on foot
Antipredator behaviour in fallow deer
145
When analysing the fawns’ and mothers’ behaviour in the main biotopes
(marshes, rush areas and meadows) the tendency was similar; so, the flight probability
by fawns younger than 4 days was significantly low (*=0.12; SD=0.12; x
5.44;
P<0.05, yJ = 14.74; P<O.O1 and Ic = 12; Pc 0.01 repectively); the mother’s flight
tendency was high in the three biotopes (* = 0.74; SD = 0.36) but due to the small
11; P<o.01).
sample it was significant only for rush areas (x?
Finally, counting on 13 cases of fawns ohserved daily, the average date of
integration into the group was calculated. The criterion being the age of the young
when seen in a group for the second time. This value was of 13.61 days (±4.31).
DISCUSSION
Most of fallow deer births take place in Doñana during the first fortnight of
June. In this characteristic the population coincides with most deer populations in
temperate areas presenting a reproductive period which is markedly seasonal and
extraordinarily synchronized (Vos 1960, MITCHELL & LINCOLN 1973, BERGERUD
1975, CLUTTON-BROCK & GUINNESS 1975).
Two general hypothesis are invoked to explain birth synchrony: (i) optimal
timing with respect to season may enhance the survival and growth of offspring as
well as the survival and future reproductive success of the mother (DAUPHINE &
MCCLURE 1974, BUNNELL 1980, DUNBAR 1980, CLUTTON-BROCIC et al. 1982, RUTBERG 1987) and (ii) females may synchronize births in order to reduce predation on
newborns either by satiating or confusing predators (DAUPHINE & MCCLURE 1974,
EsTEs & ESTES 1979, LOTT 1981, BERGERUD et a!. 1984).
With respect to predators, as already mentioned in the Introduction, at present
there are no large predators in the area and there is only the lynx (Lynx pardina) to be
mentioned, which attacks young individuals in autumn and winter, affecting mortality
rates to a minimum degree (DELIBES 1980, BELTRAN et al. 1985). However, until the
forties there were wolves (Canis lupus) living in Doñana; so, predation was an
important force selecting for breeding synchrony in fallow deer.
Furthermore, the cycle of forage availability, in an area like Donana with a very
short spring and a very dry summer, is also probably an important factor selecting the
time of the calving season. The fact that none of the offspring of late breeding
survived the summer seems to confirm this hypothesis. As in other ungulate species
(i.e. FESTA-BIANCHET 1988) inadequate nutrition is suggested as the cause of mortality
of late newborn calves.
At Doñana a peak of births has been detected coinciding with the period of
minimal diurnal activity of predators in the area (DELIBES 1980, ALVAREZ et al. 1983,
PALOMARES 1986, BELTRAN 1988, RAU 1988). A similar diurnal birth distribution has
been described for other fallow deer populations (CHAPMAN & CHAPMAN 1975,
STERBA & KLOSAK 1984) and for other ungulate species (E5TEs 1966, GOSLING 1969).
ESTES (1966) suggests that the morning peak allows the calves to gain coordination
before night fall, when predators are more active, but, since we have no data from
nocturnal observations, in our case it is not possible hypothesize.
The rapid removal of the placenta by placentophagia also lessens the possibility
that predators will be attracted to the slowly-developing calf. In some species a
diurnal distribution of breeding has been interpreted as a possible thermal regulation
C. San José and F. Braza
146
(Len 1981) but we think that this could be less adaptative in Doñana where the
mean minimal temperature itt May-June never falls bellow 10 OC in June
(i=z12.51*1.72 °C during the period 1983-1986).
The efficiency of *clying-outn as an antipredator behaviour is increased by the
initial cleaning of the fawn and by the mother consuming his faeces. These factors
reduce the chance of predator detecting a calf by its smell. This behaviour has been
reported for many ungulate species (GOSLING 1969, ESPMARK 1971, JACKSON et al.
1972, KITCHEN 1974, AUTENRIETH & FICHTER 1975, GUBERNIK 1980, TRILLMICH
1981, SADLEIR 1984).
On the basis of behavioural aspects of antipredator tactics of fallow deer during
the calving season, these species could be classified as <<hider since the newborn lie
concealed for their first few days of life. Accompanied by the concealment of young
calves, the isolation of the female reduces the chance of predation by making
detection by predators more difficult. This behaviour is usually shown by species that
calve on shrublands which vegetation provides a good concealment (ALTMANN 1963,
HAWKINS & KLIMSTRA 1970, JACKSoN et al. 1972, Wmm et al. 1972, HEIDEMANN
1973, MEIER 1973, CHAPMAN & CHAPMAN 1975, CLUTTON-BROCK & GUINNESS 1975,
NELSON & MEal 1981, OZOGA & VERME 1986). Females of species such as wildebeest
that calve on short-grass areas which offer less conceal, do not seek isolation for
parturition and there is no lying-out behaviour (ESTES 1966). In Doñana, fallow deer
have adapted a <<hiden> strategy to an open habitat, selecting the rush areas as the
biotope that offers the highest cover, and then displaying a typical thiders. behaviour
as shown by other fallow deer living in forests (GanERT 1968, CHAPMAN & CHAPMAN
1973).
On the other hand, the cryptic colour of the fallow deer newborn calves and the
particular position that they adopt lying-out could be interpreted as a complete
strategy of camouflage (TINBERGEN et al. 1967) and permits an understanding of the
motionless behaviour displayed in the open biotopes (marshes and meadows).
Furthermore, the success of the <hider, strategy in ungulates depends in part on
the mother’s habitity to minimize the information she transmits about her young’s
hiding place while remaining close enough to distract or drive away a predator
(ALTMANN 1963, LANGMAN 1977, MACCONNELL-YOUNT & SMITH 1978, PRATT &
ANDERSON 1979, TRUE’rr 1979, STEIGERS & FLINDERS 1981, BYERS & BYERS 1983).
At the approach of a potential predator (a person on foot) most fallow deer mothers
run away from the place where the young is hidden, attracting the attention of the
predator by their flight.
Finally, the effectiveness of lying-out in a very young calf depends on a flight
distance of almost zero (TINBERGEN et al, 1967, GOSLKNG 1969). A predator may then
pass close by without the calf revealing its presence by flight. In contrast, a few days
later, fallow deer calves jump and run away when approached to within a few metres.
This is probably correlated with improved chance of escape from predators by flight.
ACKNOWLEDGEMENTS
This study was supported by the Comislén Ioterininisterial de Ciencias
and the Junta de Andalucia.
y
TecnologIa (C1CYT)
Antipredator behaviour in fallow deer
147
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