George A. Bubeník Anthony B. Bubeník
Horns, Pronghorns, and Antlers
Evolution, Morphology, Physiology,
and Social Significance___________
© 1990 Springer-Verlag New York, Inc.
Printed in the United States of America.
New York Berlin Heidelberg
London Paris Tokyo Hong Kong
Social Status and Antler
Development in Red Deer
Close correlations between social dominance and levels of some hormones modu-
lated mainly by agonistic behavior have been reported in mammals. The hormone
changes which accompany agonistic interactions appear to be more dramatic and
longer lasting than those associated, for example, with sexual interactions (Hard
ing 1981). Clearly, dominant animals generally have lower pituitary/adrenocorti
cal activities than submissive animals living with them. Thus, the dominant
position usually based on aggressive behavior often tends to be related to elevated
androgen level, while subordinate status seems to be associated with lower andro-
gen secretion and increasing levels of glucocorticoids (Brain 1980; Leshner 1980).
Increased chronic ACTH/glucocorticoid production in subordinate animals sup-
presses androgens (Brain 1980). In Cervidae the evidence for these relationships
has been obtained in whitetailed deer (Bubenik, A. & Bubenk, G. 1976b; Forand
et al. 1985), red deer (Short 1979), and reindeer (Stokkan et al. 1980).
In theory, the mentioned hormones may be involved in both antler cycle timing
and antler growth. In antlerogenesis the main role seems to be played by andro-
gens. Typically, the more masculine a mammal male appears, the higher testoster-
one concentration in his blood (Crenshaw 1983). Deer are not an exception. Since
the beginning of antler growth, androgens are probably the leading hormones
affecting this process (Bubenik, G. 1982). Crenshaw (1983) injected GnRH into
immature male whitetailed deer and then determined testosterone release. He
obtained high correlation coefficients between testosterone response ratio and
various antler measurements. In another experiment, Brown et al. (1978) found
positive correlations between increasing serum androgen concentrations and
increasing antler mass in bucks. Contrary to physiological levels of androgens,
glucocorticoids were found to suppress antler growth (Bubenik, A. et al. 1976).
The above brief review allows one to put forward the hypotheses that domi-
nance in a male deer is related to: (a) the timing of his antler cycle, i.e., dates of
antler casting and/or cleaning; and (b) the process of his antler growth.
17. Social Status and Antler Development 443
FIGURE 1. Number of deer in the observed population. Open circles = hinds. Full circles
To test these hypotheses, we studied an enclosed population of "white" red
deer. The following text is based on published results of the study with some
recent data added.
Red Deer Population Studied
The study was carried out in the Žehušice Game Reserve, Central Bohemia,
Czechoslovakia, a fenced park of 2.42 square km, divided into two enclosures.
In this report we used records from the population kept in the main enclosure
(1.26 square km).
The subjects were male members of a herd of red deer, Cervus elaphus, con-
taining many white and partially colored individuals (Bartoš 1980). No red deer
in the reserve are culled, except for wild-colored male progeny. Fig. 1 shows the
number of hinds and stags present in the main enclosure over the study period.
From birth, all stags are identified individually from coat color and physical vari-
ation, so their ages are known exactly. Our analysis has involved all stags older
than 2 years.
Observations were made between 1972 and 1985 (once every 4 weeks), except
during the time of antler casting or cleaning, when they were made daily. The
animals were fed almost every day during the whole year, and so could be inspected
by an observer seated on a tractor during feeding at a distance of approximately
444 L. Bartoš
20 m without any apparent disturbance. The records of stag dominance hierarchy
were made at the time of feeding. The observations of feeding deer lasted 10-70
min, until the animals left the feeding area. During observations, the food was
always deposited in one place to induce competition among the stags. All animals
encountered each other regularly, and if one animal moved away when approached
by another, this was taken as an indication of subordinance. The outcome of such
encounters was invariably clear. The rank order was based on the encounters of
single stags with each animal of the bachelor group. The dates of antler casting and
cleaning were recorded daily by the deerkeeper.
For the analysis, several relative values are used: dominance index (DI) (Bartoš
& Hyánek 1982a)—calculated so that the position in a hierarchy (alpha=l, etc.)
was divided by the number of males present. In those parts of the study, where
it was necessary to study social position of stags in detail, DI is expressed in two
forms (Bartoš & Perner 1985): general dominance index (GDI) —the position in
the hierarchy of all the stags living within the same enclosure divided by their
total number. When used for a single observation of social groups, a measure
called relative dominance index (RDI) was used, i.e., DI within the group being
Once the rut was over, stags were separated temporarily from the main herd of
mer, toward the period of antler cleaning. Although the mean age of the stags
monitored was rather low, the alpha stags were not usually the oldest ones (Bartoš
1986a). There were very stable social relationships among the stags throughout
the year in our study herd. Changes in the stag dominance hierarchy increased
with the number of individuals present. The frequency of rank changes during the
period with hard antlers was significantly higher than that during the velvet
period (Bartoš 1986b). Between 1972 and 1983 stags developed a typical linear
hierarchy. Triangular relationships occurred seldomly, being usually a temporal
result of rare changes in the hierarchy during the period of antler casting. Only
during and after the antler casting of 1984 the stability of the hierarchy decreased
and the first permanent triangular relationships appeared. By the end of the year
the hierarchy was stabilized again, this time with numerous nonlinear dominance
relationships (Fig. 2).
Increasing the size of a social group affected animals at the two extremes of the
hierarchy (the alphas and omegas). Increasing group size elicited an increase in
agonistic activity in the former and suppressed it in the latter. The stags which
occupied the middle range of the hierarchy showed a decrease in number of
agonistic interactions when group size increased (Bartoš 1986a).
Structure of dominance hierarchies in three different seasons related to calcu-
lated rank, date of antler casting, and ages of stags.
17. Social Status and Antler Development 445
446 L. Bartoš
TABLE 1. Partial correlation coefficients between domi-
nance and antlertesting dates, and between dominance
index and antlercleaning dates in individual seasons (with
*According to Anděl (1978), the hypothesis was tested that the cor-
relation coefficients are equal. The hypothesis could not be
rejected (P = 0.05) for both antler casting and cleaning—hence,
the estimation using z transformation for all seasons was made.
Social Status and Antler Cycle Timing
The first step of the investigation was to relate rank position to dates of antler
casting and cleaning. The initial study (Bartoš 1980) indicated that the antler
casting time of individual stags was dependent primarily on social status and that
the influence of age was of secondary importance. The stags of higher rank also
tended to shed velvet earlier. After a longer period of time, partial correlation
coefficients were calculated for each season of the period when the stags estab
lished linear hierarchy in Žehušice (x = DI, y = date of antler casting/cleaning,
z = age; Table 1). (Incidental asynchronous cast was calculated as a mean date
of casting of the left and right antlers.)
Both relationships (between DI and casting/cleaning) when influence of age
was eliminated, reached high statistically significant values. The first hypothesis
advanced has been confirmed. The next step was to estimate social factors which
could influence the relationships.
Social Structure and Antler Cycle Timing
The relationships between antler casting and social position under the situation
of stabilized linear and nonlinear hierarchy were compared according to non
linear social relationships occurring since the fall of 1984. For this purpose we
used the data from 1982 (typical linear hierarchy), 1983 (linear hierarchy with an
unusually young stag in the alpha position), and 1985 (nonlinear hierarchy; see
Fig. 2). Data for spikers (deer with first antlers) were not included. For the pur
17. Social Status and Antler Development 447
TABLE 2. Comparison of correlations existing within different types of hierarchy
pose of this part of the study, the hierarchy of 1985 was estimated according to
Clutton-Brock et al. (1982). The values were ranked. Order represented a rank
position ofa stag ("rank"). The bachelor groups of stags of 1982,1983, and 1985,
respectively, did not differ in mean ( SE), dates of antler casting (March 31
5 days, March 31 4 days, March 24 4 days, ANOVA, F(2,37) = 0.90, NS),
or in mean ages (4.92 0.53,5.21 0.60,6.00 0.55 years, ANOVA, F(2,37)
= 0.98, NS). The data for each season were calculated separately using partial
correlations (x = DI, y = date of casting, z = age of the stag). The results are
shown in Table 2.
In seasons with linear hierarchy (1982, 1983), correlations between rank and
date of antler casting (with standardized age) reached high, significant values,
while in the season with nonlinear hierarchy it did not show such a close relation-
ship. Thus, the linearity of a hierarchy seems to be one of the important factors
that allows a close relationship between social position of a stag and his antler-
Agonistic Activity and Antler Cycle Timing
Years lasting observations allowed more general analysis using data of cycle tim-
ing and social characteristics per unit calendar year. It was found that the
strength of the relationship between stag rank and casting and cleaning order
under the situation of stabilized hierarchy was significantly correlated with most
of the indicators of general aggression (such as the number of killed stags, inci-
dence of broken antlers, etc.; Bartoš 1986b). The higher the level of aggression
within the herd of stags, the closer the relationship between rank and the timing
of the antler cycle indicating that the process of antler cycle timing can be modi-
fied by an aggressive behavior of a stag related to his rank. To prove directly this
suggestion, agonistic activity of selected stags was monitored in detail during
competition at feeding before casting and before cleaning. Animals of the top,
middle, and bottom rank were included. There were significant correlations
between the casting time and the most frequent aggressive acts prior to casting.
The more aggressive a stag, the earlier the date of casting. On the other hand,
there were no statistically significant correlations between agonistic activities
recorded during a velvet period and the date of antler cleaning (Bartoš 1985).
This is consistent with the results presented in Table 1. It can be seen there that
448 L. Bartoš
although both antler casting and cleaning correlated highly with DI, the correla-
tion of the former was markedly higher than that of the latter. Is there any differ-
ence between the periods preceding the casting and cleaning?
Before antler casting (i.e., since the end of the rut), the stags of the studied
herd lived usually in one large group, whereas after casting they tend to disperse
into numerous unstable small groups not separated exclusively from a company
of hinds. Therefore, the stags may be in a different social environment in the two
periods. Presumably, in a situation of changing sizes and memberships of groups,
animals of lower rank may increase their rank temporarily if they are in a group
of the lowest-ranking individuals. On the other hand, dominant animals sepa-
rated from the company of others need not have sufficient social stimulation
which would influence their internal environment. If it is so, then sample
monitoring of individual stags at that time could thus hardly record the same
facets of their aggressive activity. To solve the problem we conducted detailed
observations of the composition of individual social groups of stags throughout
the velvet period (Bartoš & Perner 1985). That is, from the time just after antler
casting of all stags to the time when the last stag is cleaned. The observations
were made approximately every other day within the velvet period. As in other
red deer populations (Appleby 1983; Bützler 1974; Clutton-Brock et al. 1982;
Darling 1937), our stags tended to associate with animals of similar rank and age.
RDI values were calculated for each stag for each observation and were compared
with antler cleaning dates. The correlation coefficients increased toward the time
of cleaning. For the last 2 weeks of the period the coefficients reached levels simi-
lar to those between stags' rank and antler casting. To complete the analysis, the
association between stags was defined. The closest associates of each stag (deter-
mined after Appleby 1983) in the weeks preceding each individual's antler clean-
ing date were identified. The higher-ranked associates of the same age cleaned
significantly earlier than the lower-ranked individual. It was concluded that both
antler casting and cleaning are regulated by hormones modulated by agonistic
behavior related to rank (Bartoš & Perner 1985).
Antler Casting in Different Cervid Species
Until now we have discussed the relationships between rank position and antler
cycle timing in red deer. Forand et al. (1985), observing captive herds of white-
tailed deer, found a highly significant inverse correlation between rank and order
of casting antlers, indicating that dominant white-tailed deer bucks retained their
antlers longer than subordinates. This result is in apparent contrast to what has
been found in our red deer herd (Bartoš 1980, 1986b). The authors (Forand et al.
1985) presented a brief review giving literary evidence that in northern areas of
the United States where antler casting is early and relatively short (from mid-
December and to late January), older, larger, and presumably dominant bucks
cast antlers earlier than their subordinates. For the Midwest, where antler cast-
ing extends from January to late March, white-tailed bucks with large antlers
generally retain them longer than bucks with small antlers. The authors sug-
17. Social Status and Antler Development 449
gested that dominant whitetails in northern ranges may experience considerable
stress and physical exhaustion due to a short but intensive rutting season. Then
they related the suggested stress with increased levels of corticoids reducing
levels of testosterone. They concluded that testosterone levels of subordinate
animals may stay longer above the threshold for antler casting, resulting in longer
retention of antlers.
Are the presented contradictory results based on a methodological mistake, or is
there any speciesspecific differences? The latter seems to be more likely. It is our
suggestion that the difference between red deer and whitetails in the relationship
between rank position and the order in which individuals cast their antlers lies in
the speciesspecific response of antler casting to seasonal pattern of testosterone.
Brown et al. (1983a) have shown that the endocrine control of the antler cycles of
two deer species may differ. It has been well documented that there are species
specific differences in seasonal patterns of casting and new antler growth. Deer
species with seasonally determined antler cycles may be divided into two basic
groups. Group A—those in which casting of old antlers is followed immediately by
growing of a new antler, such as red deer, wapiti, sika, fallow, roe, and Pere David's
deer; Group B—those in which an interval exists between antler casting and new
antler growth such as whitetailed deer, muledeer, moose, reindeer, and caribou
(Bubenik, A. 1966; Goss 1983; Jaczewski 1981a; Sempéré & Boissin 1982). In red
deer, antler casting and regrowth are closely interdependent. Our hypothesis about
the positive correlation between antler casting and rank position of a male fits well
(Bartoš 1980, 1986b). It is presumed that this might be the case for all species
belonging to Group A. On the other hand, in species in Group B, antler casting and
starting of new antler growth are well separated in time, so that they may have dif-
ferent relationships to rank position. While high rank position may associate with
the delay of antler casting, as has been found by Forand et al. (1985), starting of
new antler growth should be enhanced. In other words, the time between antler
casting and new antler growth should be shortest in the highestranking animals of
Group B. To support the above hypothesis, we can submit some empirical data. For
species in Group A: in red deer (Bützler 1974; Lincoln 1972; Lydekker 1898;
Nečas 1959), and wapiti (Bubenik A. 1982a), it is well established that the older
and stronger males cast earlier. For species in Group B: Kozhukchov (1973)
reported in farmed bull moose that young animals cast antlers 13 months earlier
than old animals.
How can we explain the relationships between the behavior and antler casting
and the difference between the two groups of deer species on a hormonal basis?
While males of the species belonging to the Group A need some stimulation of a
new antler growth, it is the decline of testosterone itself after the rutting season
that seems to be responsible for the casting of the antlers which may occur in early
winter in males of Group B (Brown et al. 1983a; Mirarchi et al. 1977b). White
tailed bucks treated with antiandrogen cyproterone acetate immediately after
casting their old antlers did not renew growth of antlers (Bubenik, G. 1982). It
can be predicted that the treatment of males in Group A with antiandrogen before
casting might respond in retaining their antlers. Red deer stags that die in spring
450 L. Bartoš
An overaged stag (in the foreground) who failed to cast antlers in the spring is
sparring with a mature stag carrying fully developed velvet antlers.
in Scotland are almost invariably still carrying their hard antlers (Mitchel in Lin-
coln 1971a; Lincoln & Bubenik, G. 1985). In 1985, the oldest stag of the
Žehušice herd failed to cast his antlers (Fig. 3). He had retained them until next
October, when unfortunately he was shot. It was found that he had fully regressed
testes. The evident fall of testosterone levels was not sufficient to induce casting.
On the other hand, van Ballenberghe (1982) reported several cases of bull moose
which were handicapped by an injury and which cast earlier than others, suggest-
17. Social Status and Antler Development 451
ing that the fall of testosterone levels was potent enough to initiate casting.
The presumed speciesspecific response to seasonal testosterone variation may
be reflected also in the cast antlers. The longer dead antler is attached to the
pedicle, the more it dies back (Lincoln 1984). The species which cast soon after
the rut tend to have a convex casting surface at the base of the cast antlers com-
pared with the concave casting surface in the species which cast later. This is also
apparent when red deer are castrated soon after the rut (Lincoln 1984; Lincoln &
It has been hypothesized that new antler growth may be initiated by a small
reactivation of sexual function and hence testosterone pulse (Bartoš 1980;
Bubenik, G. 1982; Goss 1983; Sempéré & Boissin 1982). This may correspond
to the initiation of pedicle formation within a male's ontogeny. It has been shown
that during puberty the deer testes must be activated for a short time to induce the
growth of pedicles (Brown etal. 1983a, 1983b; Lincoln 1971; Sempéré & Boissin
1982; Suttie et al. 1984). The initial suggestion that new antler growth may be
induced by the shortterm pulse of testosterone was based more or less on indirect
results (Bartoš 1980). However, now there are rather more direct indications
available. Testosterone titers were determined to increase twice a year in red deer
(Blaxter et al. 1974; Suttie et al. 1984), wapiti (Haigh et al. 1984), whitetailed
deer (Bubenik, A. 1984; Bubenik, G. et al. 1982a; Brown et al. 1983b; Mirarchi
et al. 1977b), and sika deer (Brown et al. 1983a). The effect of testosterone,
modulated by behavior, on the initiation of antler regrowth could probably be
determined by measuring the absolute hormone level. Low amounts of testoster-
one can stimulate bone growth, and higher levels may be inhibiting (Brown et al.
1978b). The more dominant males have earlier, higher, and more frequent
testosterone pulses within the range of Brown et al.'s 'low amount'; a new antler
bone growth may thus be initiated more vigorously and the antler casting in spe-
cies in Group A may occur earlier (Bartoš 1980). In males in Group B, new
antler growth of dominants may start earlier even though antler casting had
In general, fighting stimulates the level of glucocorticoids (see Introduction).
Increased corticoid levels inhibit antler growth (Bubenik, A. et al. 1976).
Adrenal hypertrophy may occur in deer under conditions of stress (Bubenik, G.
& Bubenik, A. 1965; Hughes and Mall 1958), a reaction which may delay antler
casting in fallow and red deer (Fig. 4) (Bubenik, G. 1982; Topiňski 1975),
representatives of species Group A. Hypothetically, the species of Group B
should be affected in just the opposite way by stress, enhancing antler casting. A
role of some other hormones may be expected. Nevertheless, this possibility is
not included in this simplified model.
If our earlier hypothesis is correct, how does it fit to antler casting caused by
castration? From the point of view of the hypothesis, we should expect a sudden
fall in testosterone levels after castration, followed by temporary restoration of
androgen levels (of adrenal origin?) which afterward definitely declines. To our
knowledge, the detailed pattern of testosterone decline after castration has not
been investigated. However, there are some indirect data. After orchidectomy,
452 L. Bartoš
FIGURE 4. Asynchronously casting stags are on average older and higher ranking than
those casting synchronously. Among the asynchronously casting stags, the higher a stag
ranked, the shorter interval between the dates of casting of both his antlers (Bartoš and
17. Social Status and Antler Development 453
LH levels increased more than four times in red deer (Lincoln & Kay 1979) and
more than four times in white-tailed bucks (Bubeník, G. et al. 1982a). In theory,
testosterone secretion from the adrenal gland may be facilitated by this way to a
short-term surge. Under such circumstances the difference between the two
groups of Cervids should be diminished. This seems to be the case. As reported
by Goss (1983), after castration, renewed antler growth occurs soon after the old
antlers have been lost, even in those species in which there is normally a lag
between these two events. However, the time of antler regrowth after castration
is season-dependent in both groups of the deer species (Lincoln 1984; Bubenik,
G., personal communication).
How would the above hypothesis explain that some solitary-living old individ-
uals may cast their antlers rather early in the period of antler casting when com-
pared to most of other conspecifics in Zehuiice while performing minimal social
interactions? In experiments with artificially altered daylight and its influence on
the antler cycle, it was shown that older males can sometimes express endoge-
nous annual antler growth, irrespective of artificial light conditions (Goss
1969a). This may be the case in solitary-living stags. On the other hand, the
absence of social stimulation may cause the observed fact that these solitary-
living stags always cast antlers later than the younger top dominant ones living in
social groups (Bartoš 1980).
It must be emphasized, however, that general good nutritional status of a popu-
lation seems to be an essential factor allowing an expression of the behavior in
antlerogenesis. Suttie (1980a) found in red deer that the quality of nutrition
influences the seasonal levels of testosterone and prolactin. Good nutrition
caused even rutting of his stags twice a year, in spring and fall. In this respect, it
is important to note that there is also sufficient evidence confirming a species-
specific dependence of antler casting on nutrition. In red deer, Darling (1937)
and A. Bubenik (1966) argued that inadequate nutrition after the rut significantly
delays casting. Also, Lincoln (1971a) stated that red deer stags in poor condition
cast their antlers later. Watson (1971) showed that antler casting among red deer
was delayed by food restriction brought about by severe weather conditions, but
supplemental feeding could reverse this trend. Similarly, in experiments with
farmed deer, Suttie & Kay (1982) and Fennessy & Suttie (1985) found a trend for
antlers of a nutritionally unrestricted group to be cast 1-2 weeks before the res-
tricted group. In contrast, Long et al. (1959) showed that nutritional deprivation
of white-tailed deer in the spring hastened antler casting. Also Lincoln & G.
Bubenik (1985) stated that white-tailed deer cast their antlers earlier than normal
in winter in response to poor feeding and loss of condition, while the regrowth
of new antlers in such animals occurs later than normal in spring. Ozoga & Verme
(1982) reported an influence of improved nutrition on antler casting in white-
tailed deer. Supplementally fed bucks in the enclosure retained their antlers
several months longer than did the bucks in that area. West & Nordan (1976a)
found similar relations in mule-deer. Better fed captive bucks often cast antlers
later than wild bucks.
454 L. Bartoš
Conversely some other data do not support the above hypothesis, either for
Group A or Group B. Gibson & Guinness (1980) reported, for example, that red
deer stags with the highest reproductive success (and hence of the highest rank)
during the rut cast their antlers later than others. Presumably poor condition nega-
tively affects the spring testosterone pulsation of these animals as in Suttie's
(1980a) restricted group [see also the abovementioned experience of Darling
(1937) and A. Bubenik (1966)]. The same explanation may account for all the dis-
crepancies in casting of whitetailed deer of various geographical origin cited
earlier (Forand et al. 1985). In northern areas, one may expect worse nutrition of
the deer than in southern areas. Dominant bucks are not able to maintain levels of
testosterone above the threshold for antler casting because they are significantly
more exhausted after rutting activity than their subordinates. Earlier casting in
dominant bucks thus may occur. Moreover, seasonal fall in testosterone levels
under conditions of naturally restricted nutrition cannot be stimulated substan-
tially by dominant related behavior, which may also explain why the casting time
in whitetailed deer is more synchronized and occurs earlier in the season in north-
ern than in southern areas as reported by Forand et al. (1985). Under better general
conditions, in the Midwest, such behavior may lead to maintenance of elevated
testosterone levels of dominant bucks causing the delay of antler casting.
Antler Cleaning and Deer Species
Contrary to casting, antler cleaning seems to follow the same pattern in both
groups of deer species in relation to social position. A tendency for a positive
correlation between rank and order in which males clean antlers was found not
only in our red deer (Table 1), but also in captive whitetailed deer (Forand et al.
1985). The stimulatory effects of social interactions among dominant males
probably elevate levels of testosterone, while the interactions elevate glucocorti-
coids and depress testosterone levels in subordinates. As a result, antler cleaning
may occur earlier in dominants and later in subordinates.
Many authors have suggested that antler cleaning dates are fully dependent on
age, such as in red deer (Butzler 1974; Darling 1937; Nečas 1959) and in other
cervids (e.g., Hirth 1977). Spikeantlerěd deer clean antlers later than fork
antlered males in red deer (Bubenik, A. 1966; Darling 1937; Jaczewski 1981a;
Lincoln 1971b; Nečas 1959), whitetailed deer (Hirth 1977; Jacobson & Griffin
1982; Scanlon 1977), and moose (van Ballenberghe 1982). Nevertheless, there
are also contradictory reports, mainly from captive populations. Both the earliest
and the latest cleaning dates were observed among yearlings in fallow deer
(Chapman & Chapman 1975; Štěrba & Klusák 1984), in whitetailed deer
(Jacobson & Griffin 1982), and in our red deer herd. Exceptionally early cleaning
by yearlings may be observed under natural conditions, too [e.g., in moose (van
Ballenberghe 1982)]. Here again, different opportunities for social grouping and
hence for differential social stimulation of the process, as well as differential
opportunity to be stressed, may be involved.
17. Social Status and Antler Development 455
Social Status and Antler Growth
The possibility of a relationship between social position and antler size in Cer-
vids has been widely discussed.
Some authors have suggested that antlers advertise an individual's dominance
status (Beninde 1937; Bubenik, A. 1968, 1982b; Geist 1966b; Henshaw 1969),
but variable results have been obtained in field studies designed to assess this.
Espmark (1964) in reindeer and Lincoln (1972) in red deer found that after the
loss of antlers, either naturally or artificially, individuals became less effective in
competition with other males, resulting in loss of social rank in the bachelor
group. Biitzler (1974) measured the lengths and weights of cast antlers from stags
of known social position. There was a positive correlation between these mea-
surements and social position but the author questioned whether the relationship
was genuine. Clutton-Brock et al. (1979) observed more than 100 rutting fights
between red deer on the isle of Rhum in Scotland and found a weak correlation
between the number of points on the antlers and fighting ability. No relationship
between antler length and fighting success was apparent. Appleby (1982) also
observed red deer of the same population and found that the rank of mature stags
in winter was correlated to antler length. Winter rank in mature stags was, how-
ever, correlated significantly with the weight of their antlers in one of two study
years. Suttie (1980b) found in a group of farmed stags that antler weight but not
antler length or number of points was positively correlated with social position.
Miura (1984) reported for male sika deer that large antlers were related to
All the mentioned studies were based on observations at the time when the
male deer had completed their antler growth. That is, the studies compared the
relationship between males' fighting abilities and size of their grown antlers. The
criticism of the suggested behavioral significance of grown antlers was made in
red deer during the rutting season. The fighting ability of individual stags
changes during the course of the rut as their body condition declines, and
individuals vary in the timing of their declines. Consequently, a stag that assessed
its opponents on criteria that did not vary with changes in body condition during
the rut would take many incorrect decisions (Clutton-Brock et al. 1979, 1982).
In our previous studies we have already suggested that social position and related
agonistic activity of stags during the velvet period influence the antler weight,
length and number of points, and therefore the size of grown antlers are a conse-
quence of previous social position and not vice versa (Bartoš & Hyánek 1982a,
1982b). Also Wólfel (1983) claimed possible role of rank position of a male
yearling red deer in antler development. The aim of the following study was to
assess in detail how the social position of a growing red deer stag is related to var-
ious measures of antler development.
In red deer, the gain in antler growth or development correlates in the first 5-6
years almost linearly with body growth and weight. Afterwards, i.e., after the
stags have reached their mature body size, there is always a substantial variation
456 L. Bartoš
TABLE 3. Relationships of general and relative dominance indices to antler weight and
length, and number of antler points
General dominance index (GDI)
Relative dominance index (RDI)
No. of antler points
(Bubenik, A. 1966, 1982; Huxley 1931). Hence we have simplified the antler
growth into a model that states that antlers increase linearly during a stag's
ontogeny up to 5 years of his age, while afterwards there is not regular increase.
[The same pattern has been found also in a red deer stag's social position by
Appleby (1980) and Bartoš & Hyánek (1982a).] According to the model pro-
posed, a linear regression of growth of all the antler measurements and
dominance indices (DI's) during the period of antler growth was estimated for
each stag. Then values for two extreme ages of a stag's ontogeny were calculated
(2 years—the beginning of the first regular branched antler growth, and 5
years —the end of body development). Such calculated values of the DI were cor-
related with those of antler measurements. The results of the analysis (Bartoš
et al. 1988) showed that high-ranking stags of both extreme ages had heavier,
longer and more branched antlers. The social position involved (DI) had always
been estimated during the period of formation of the future antler. However, as
mentioned earlier, in Zehušice the bachelor group tended to disintegrate during
the velvet period. Thus DI does not reflect detailed changes in social environ-
ment. Hence we further presumed: If we compared GDI and RDI throughout the
period of antler growth of a stag, then correlation between antler size characteris-
tics and the indices should fit better to RDI than to former one. To solve the
problem we used the data of 1983 when we followed the distribution of all stags
in individual social groups. We used the records of the period between antler
casting and cleaning for each stag. It represented 59.71 ±2.70 different observa-
tions for a stag. Cast antlers of 11 individuals were collected during the following
spring and measured. The values of GDI of the velvet period of these stags were
equal to those of RDI in one case, while they were not equal in ten cases (Sign
test, P = 0.01). All the values used in the analysis (GDI, RDI, antler weight and
length, and number of points) were adjusted for age using an analysis of linear
regression (Snedecor & Cochrane 1965). Correlation coefficients between the
indices and selected antler size characteristics are shown in Table 3.
The correlation coefficients between antler characteristics and GDI were
lower than those between antler characteristics and RDI in all three cases. So the
presumption has been fully supported.
The results presented should be taken as applying to a model. The living condi-
tions in Zehušice differ in several ways from those elsewhere. The situation in
17. Social Status and Antler Development 457
mature stag who cast one antler is "flailing" in an attempt to hold the rank
over a subadult, antler-carrying stag.
this herd is difficult to compare with the free-living individuals. The deer could
not leave the fenced area. So that the animals were unable to separate fully from
others and interactions with conspecifics could not be avoided. From Suttie's
(1985) observations of farmed red deer stags, it was apparent that although a sta-
ble hierarchy existed in bachelor herds, it did not prevent aggression. Suttie
found that the level of aggression was much higher than in the wild. He concluded
that members of a hierarchy may be stressed to an extent not seen in the wild.
Hence physiological consequences of aggression related to rank position under
conditions of restricted space may be expressed more markedly than those under
natural conditions. On the other hand, it seems to be likely that a population such
as ours may be a relevant model for free-living populations, since the conse-
quences of high density in the pen which may influence antler structure (and/or
antler cycle timing) are more evident under these defined living conditions. The
behavioral significance of the antlers may be of secondary importance depending
on social background of a studied herd and/or on previous social experience of
a male deer. In Žehušice, the bachelor group of stags is constant throughout the
year. The same animals encounter each other during the velvet period as during
the rest of the year. This results in a very stable social hierarchy. Social status
remains almost constant even after antler casting (Fig. 5) or antler breakage
(Bartoš 1986b). It is no wonder that there is a close correlation between rank
position and antler length, antler weight and the number of tines found also out
458 L. Bartoš
of the velvet period (Bartoš & Hyánek 1982b). In freeliving populations, reduc-
tion in gregariousness immediately after antler casting is known among red deer
stags (Biitzler 1974; Geist 1982; Lincoln et al. 1970). This may result in a less
pronounced relationship between rank position and antler size. Just before the
rut, bachelor groups completely disintegrate and stags widely disperse (Bützler
1974; CluttonBrock et al. 1982; Darling 1937; Lincoln etal. 1970; Nečas 1959).
During the rutting season, strange stags may be encountered. Sexual competition
during that time brings a strong motivation for fighting. This may encourage
stags to ignore experience gained in a bachelor group during the antler growth
that taught them to avoid an interaction with larger antlered individuals. That is
probably why CluttonBrock et al. (1979) did not find any simple relationship
between success in rutting fights and antler size. After the rut, stags usually
return to their original ranges (Biitzler 1974; CluttonBrock et al. 1982; Darling
1937; Lincoln et al. 1970). Winter groups thus consist mainly of stags which had
been present during antler development. Hence Appleby (1982) was able to
detect at least partly some relationships between rank and antler characteristics
in his winter observations. The behavioral meaning of fully grown antlers and the
physiological consequence of the stag's behavior on antler growth seem to have
quite a different basis. Behavioral meaning of the antler size of red deer stags
probably depends on social background of the studied population and previous
experience of an individual. Hence, there are a number of studies suggesting no
consistent tendency for males to avoid fighting individuals with larger antlers, at
least in red deer (Appleby 1982; CluttonBrock et al. 1979; Krzywiňski 1978;
Lydekker 1898), while under some circumstances there is evidence of an advan-
tage to bear large antlers (Bartoš & Hyánek 1982b; Bubenk, A. 1982b). The
physiological consequence of the stag's behavior on antler growth may act since
the beginning of the velvet period. The more dominant a stag, the higher the
seasonally attained levels of androgens, the greater the enhancement of antler tis-
sue formation. This suggestion has been supported by Shilang & Shanzi (1985)
who found that a small amount of androgens to sika deer stag's food during the
velvet period stimulated their antler growth. On the other hand, the lowest rank-
ing stags may lack this androgen stimulation and the presumably elevated
glucocorticoid levels may actually suppress antler growth (see Introduction). A
fall in social hierarchy in aged stags may be an initiation of their antlers "going
back." A. Bubenik (1982a) stated that in disorganized populations, a stag could
be "over the hill" at 1011 years, as opposed to 1618 years in stags of well
organized herds. In young male deer, the size of antlers increases with increasing
body size in succeeding seasons (Bubenik A. 1966; Goss 1983; Jaczewski 1981a).
The seasonal peaks of testosterone also increase in parallel with antler and body
growth, as was shown in red deer (Bubenik, A. 1984; Lincoln 1971a), roe deer
(Sempéré & Lacroix 1982), and whitetailed deer (Bubenik & Schams 1986).
Bubenik, G. et al. (unpublished, shown in Bubenik, A. 1982a) found in red deer
that young animals had the highest levels of Cortisol which then declined in prime
stags (410 years) and then rose again in old ones (11+ years).
Cameron (1892, cited in Goss 1983) was probably the first who reported the
primary importance of body weight in a combat of deer. He said that not males
17. Social Status and Antler Development 459
with heaviest antlers are favored in fights, but those with the heaviest body
weight. Clutton-Brock (1982) has argued that antler size is related to individual
differences in body size and weight. To support it, there is a large amount of evi-
dence for a relationship between body size and weight in red deer and antler
weight (Appleby 1982; Clutton-Brock et al. 1979, 1982; Huxley 1926, 1931;
Hyvärinen et al. 1977). Initially, there was no body weight data available for our
"white" stags. However, there were some indications that body weight did not
have exclusive influence on the relationship between rank position and antler
development (Bartoš et al. 1988). In the 1985 season, we succeeded in obtaining
the data of the relative body weight of the stags under study. A close correlation
between body weight of a stag and his rank position was found when the age was
statistically standardized (r = 0.79, P<0.01). On the other hand, a nonsignifi-
cant (P>0.05) correlation was apparent between body weight of a stag and the
size of his antlers (with eliminated influence of age) under the living conditions
of the park, while rank did not correlate with several antler characteristics such
as antler length, etc. When body weight was controlled by partial correlation,
rank correlated with antler length, number of tines, number of points on the
royal, bez tine, third point of the royal, and length of all the royal points. On the
other hand, when rank was controlled by partial correlation, there was still no
significant correlation between body weight and antler characteristics (Bartoš
et al. 1988). This was contrary to results of the above-mentioned authors.
Nevertheless, it has already been observed in Scotland that live weight need not
correlate with antler size in red deer (Suttie 1980b).
It may be concluded that the relationship between rank position of a stag during
the velvet period and intensity of his antler growth does exist regardless of his
body weight. Those studies showing relationship between body weight and antler
size of stags living in groups should be analyzed also from the point of view of the
stag's rank position in the same time, since under certain conditions rank position
may be even more influential on the antler growth than the body weight.
It can be concluded that the advanced hypothesis was confirmed. Dominance
in male red deer was found to be related to timing of antler cycle and antler
growth. The dominant individuals of socially stabilized group tend to cast and/or
clean antlers earlier and produce larger antlers than subordinate ones.
Acknowledgments. I gratefully acknowledge the excellent field assistant I have
had over many years from V. Perner, the former deerkeeper at Žehušice. I am
indebted to A.B. Bubenik, G.A. Bubenik, R.N.B. Kay, and G.A. Lincoln for
stimulative criticism of an earlier draft of the whole manuscript. Helpful com-
ments of P.T. Brain, T.H. Clutton-Brock, and V. Geist on the part of this chapter
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