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The Evolution of Horn-Like Organs
Author(s): Valerius Geist

Source: Behaviour, Vol. 27, No. 3/4 (1966), pp. 175-214
Published by: BRILL
Stable URL: http://www.jstor.org/stable/4533157
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THE EVOLUTION OF HORN-LIKE ORGANS

by
VALERIUS GEIST 1)
(Zoology Department, University of British Columbia, Vancouver, B.C., Canada)
(With i6 Figures)
(Rec. I5-XII-I965)
INTRODUCTION
In the evolution of mammals horn-like organs have appeared a number
of times. We find them assuming a great variety of forms; small, skin
covered pedicles in the Okapi (Okapia), large horn curls in sheep (Ovis),
or many-pointed, bizarre bony paddles in moose (Alces). Horns originated
on different areas of the head and grew from several tissues. We find horns
on animals as distantly related as Cervus, a deer, and Mylogaulus, a miocene
rodent. Also, we find horns evolving independently a number of times in
closely related groups, such as in the ruminants and the rhinos. It is inherent
in our concept of the selective force in evolution that these organs, whatever
their shape, size or position should have been an important, adaptive feature
of their owner. Yet the evolution of horns and antlers has been a puzzling
problem. COLBERT(I955, P. 398) writes: "on the fact of it, we would think
that one good pair of horns would enable a species of antelope to protect
itself and spread widely. Yet in Africa there are literally dozens of antelope
species with an astonishing variety of horns. There are straight horned
Oryx, the twisted-horned elands, the spiral-horned Kudus, the curve-horned
sable antelopes, the recurve-horned wildebeest and so on. And many of these
antelopes seemingly live in the same environment. What, therefore, are the
advantages of a Kudu's horns over an eland's horns, or vice-versa? These
are questions still to be answered."
Since the time of COLBERT'S(I955) writing, behaviour studies on ungulates have thrown new light on the function of horns. In particular WALi) Acknowledgement: I am most grateful to Dr IAN McTAGGART-COWAN
and Dr
H. D. FISHERfor critical reading of the manuscript.
Behaviour XXVII
12
T76
VALERIUS
GEIST
.,,/
Ar
K~~~~~~~~~~~~
f'
Fig. i. Hornis evolved in great variety of form and size. Despite this divTersitythere
are somnemain functionaltypes towards which quite distantly relatcd nmammals
converged.
Such types are (I) (a) Eobasileus and (b) Protoceras. (II) (c) Tetraceros and (d)
Giraffokeryx. (TIII) (e) Arsinotheriumf,(f) Brongtopsand (g) a dicerathere. (TV) (h)
EVOLIUTION
OF HORN-LIKE
ORGANS
I77
Cerv7ss elaphus and (k) Tragelaphubs.(V) (i) Muntiacus, (1) Antilocagpraand (m)
Capreolus. The inverse relationshipofEtusks and antlers is illustrated in the series (h),
While Protoceras (b) stood at the beginning of protoceratid
(i) and (j) HIydropo3tes.
evolultion,Synthetoceras (n) was the specialised last genlus from the pliocene.
178
VALERIUS GEIST
studies have been illuminating. It has become evident that the

question of horn evolution encompasses most of the biology of horn bearers,
and cannot be treated as an independent question. In this paper, I have
attempted to consider the evolution of horns in a broad context, with special
emphasis on the functional aspect of these structures in the behaviour of
the organism.
There is no good definition for horn-like organs. As LANKESTER (1902)
points out: "The word "horn" is a comprehensive term of various and
uncertain meaning". In the restricted sense the term is properly applied
only to those structures in which cornified epidermal material plays an
important part in structure. For present purposes I would like to include all
hardened excrescences projecting from the head, no matter what their
origin. This concept of horn-like organs will include horns sensu strictu
as well as antlers and even the large tusks of suids and proboscidians. I will
therefore take an analogistic, rather than a homologistic view for reasons that
will become evident.
Horn-like organs appeared early in the history of mammals. They occurred
on the heads of Dinocerata and embrithopods (Fig. I. a, e), both families
of peanungulates from the paleocene and eocene. Among the perissodactyls
the earliest hornbearing family was that of the titanothers (Fig. I. f). The
rhinocerotids independently evolved horns at least twice. The present day
rhinos of Africa (Diceros and Ceratotherium), as well as the Asian rhinos
Rhinoceros and Dicerorhinus, probably represent the first group. The
aceratheres, hornless for almost all of their long evolutionary history, evolved
horns in their terminal genera just prior to extinction (THENIUS &
HOFER, I960). It is possible the miocene diceratheres also evolved their
horns independently. These rhinos possessed two horns side by side on the
rostrum (Fig. I. g).
In the ruminants also horn-like organs apparently arose several times
independently. The horn type of giraffids can be homologised with that of the
bovids, but not with that of the cervids. On the basis of fossil evidence it has
been proposed that the antilocaprids (including the antlered, hypsodont
Merydontidae), evolved independently, and are not closely related to
bovids (THENIUS & HOFER, I960). The protoceratids possessed horn types
that appear to be different from anything known from present day ruminants.
It is hence likely that hornlike organs originated and evolved independently
in the giraffid-bovids, the cervids, the antilocaprids and protoceratids.
In spite of the bewildering variety of horn shapes and sizes, there are
some basic structural plans which appear repeatedly. Compare the arsinothere, titanothere and dinoceratid (Fig. I. e, f, g), or the ancient dinocere
THER'S
EVOLUTION
OF HORN-LIKE
ORGANS
I 79
Eobasileus and the ruminant Protoceras (Fig. I. a, b), or Tetraceros - a

bovid - and Giraffokeryx - an extinct giraffid (Fig. I. c, d.). Since
WALTHER
(I962) suggested that the ridges and twists in antelope horns
function to hold the horns of antagonists locked together in battle, I would
propose to regard the antlers of Cervus and the horns of Tragelaphus as of
one functional type (Fig. I. h, k). The horns or antlers of Antilocapra,
Muntjacus, and Capreolus show convergent features (Fig. I. 1, i, m). In
their early evolution dinoceratids and titanotheres possessed a pair of small
horns positioned antorbitally on top of the head. In the later genera of
titanotheres the horns became anteriorly located and fused into one stem,
resembling in principal the horn type evolved later by the rhinocerotids.
Well known, but still striking, is the negative correlation of tusk and horn
size. Early rhinocerotids had permanent tusk-like lower 2nd incisors; later
genera lost these with the evolution of horns. Early genera of cervids were
antlerless but possessed large upper canines; most present day genera of
cervids while possessing large antlers, have either lost or reduced to
insignificance the upper canines. The evolutionary stages have been preserved
to the present in Hydropoltes, Muntiacus and Cervus (Fig. I. j, i, h).
Although protoceratids retained canines throughout their evolutionary history,
these were quite small in their last, large horned genera.
Not only the variety of form has been great among horn-like organs,
but also their tissue composition and basis for attachment. For illustrative
purposes the structure, histology and ontogeny of horns and antlers will be
reviewed.
The best known horns are of the giraffid-bovid and cervid type. The horns
of giraffids and bovids develop from ossicones as well as from the overlying
skin and skin outgrowths. Ossicones grow as independent bone elements
from the skin covering the forehead; they are not part of the dermal skullbone, but shortly after birth become fused to the skullbones underlying them.
In giraffes the large ossicones fuse to the parietal bones during the fourth
week of postnatal life (BRUHIN, 1953). In Okapia the ossicones are situated
on the frontals, as is the rule for bovids as well (LANKESTER, I902, GIJZEN,
I959). The ossicones grow into the horn cores of bovids or horns of giraffids.
They are porous or cancellous structures with large sinuses, and grow
rapidly during the early life of the animal but have a reduced growth rate as
the animal ages (BUBENIK,
I959). In the giraffes the skin covering the
ossicones grows a hair coat, while in the bovids this skin grows a mass of
horn. In the bovids the horn core is relatively conservative in form and it
is the overlying cornified sheath that is largely responsible for the great
variety of horn types.
I80
VALERIUS GEIST
Very little is known of the development and growth-regulating mechanisms

of bovid horns. In some bovids horns reach their definitive shape and size
early in life and do not grow appreciably after that (Gorgon, Ovibos). In
others there is a periodic annual growth throughout life (Ovis, Rupicapra).
Here horn growth is probably related to the annual cycle in sex hormone
production. The horn grows when sex hormones are at their lowest concentration and ceases to grow with the onset of the early pre-rut.
Little is known of the ontogeny of antilocaprid horns. The resemblance
to bovid horns in gross morphology is very likely spurious. One great
difference is the annual shedding of the horn sheath by Antilocapra,
whereas bovids retain their horn sheaths throughout life.
Deer antlers are true bone outgrowths from the frontals and hence are
not homologous to the os-cornus of the giraffid-bovid horn (BUBENIK,
I959). During growth, antlers are skin covered structures growing apically
(BRUHIN, 1953), and hardening by a progressive ossification of a cartilaginous substrate (BUBENIK, I959). It is probable that antler ossification takes
place at the temporary expense of the calcium phosphate of the skeleton
(MEISTER, I956). After the antlers reach definitive form, the velvet covering
driesi and is rubbed off by the animal. Sometimes it is consumed (DARLING,
I937). It has been shown that increase in weight persists for some time
after the velvet is shed, but at maturity the antlers are dead bone, cut off
from the metabolic pool (BUBENIK, I959, I963; BRUHIN, 1953), and without
sensation of their own (HECK, I956). After the rut male cervids lose the
antlers. The dehiscence involves the erosion of the bony material by osteoclastic activity at very specific regions. Antler regrowth may be immediate
as in Cervus elaphus, or after a period of dormancy as in Alces. Growth
and development of antlers is apparently differently regulated in various
species.
The rhinoceros horn is a very different organ in structure and histology.
It grows from a bony base on the nasal bone and contains no interior bony
core. The horn is formed from a dense mass of filaments growing from
dermal papillae in the skin covering the base. It was long accepted as being
formed from densely matted hair, but recent research showed that the horn
filaments are not hair. They are 300-500 microns in diameter, many-sided in
cross section, laminated in structure, centrally hollow in the core, and lacking
the cuticle so characteristic of hair. The filaments are held together by very
little interfilamentous horn. This does allow some fraying of the horn
(RYDER, I962). The horn of Indian and Java Rhinoceros species are
frequently badly worn, and have no sharp points as do those of Diceros,
Ceratotherium and Dicerorhin,cs (SODY, 1959).
I8I
EVOLUTION OF HORN-LIKE ORGANS
SOME HISTORICAL
VIEWS ON HORN FUNCTION

EVOLUTION
AND
There is no lack of opinion on horn function and evolution. BRUHIN

reviewed many of these most admirably. Therefore only some of the
more interesting views will be mentioned here. Frequently horn function
and evolution have been confused. Much evidence is of anecdotal nature,
although such evidence is very valuable to the formation of hypotheses.
The most common view is that horns evolved as weapons. LIBINSKY,
however, (BRUHIN, I953) thought antlers to function as indicators of
sexual vigor, allowing females to select the best father for their offspring.
KRIEG (BENINDE, I937) suggested that antlers evolved as metabolic valves.
According to this view, excess material from the metabolic pool would be
removed from circulation and stored in the antlers. He held the ossified antler
to be of little adaptive value. Rather, the relief from superfluous energy
and building material was adaptive. The antlers were thought to be of no
hindrance to the animal. BENINDE (I937) argues further in favour of this
view. He stated that the frontal bones were the best site for such a "'valve
organ", since here it interferes least with the essential actions of the animal.
The facts that antler growth requires much energy (BUBENIK, I959), that
ossifying antlers withdraw material from the skeleton, that deer on minimal
diets still grow antlers, that antlerless male deer do survive as effectively
as do barren female deer, all argue against the "valve" theory.
BENINDE (I937) expressed the hypothesis that antlers act as display
organs. Their greater size in vigorous, healthy adults may have an a priori
intimidating effect on the opponent, hence overt fighting is reduced since the
intimidated opponent is less likely to fight. HEDIGER (BRUHIN, I953)
expressed a similar view upon which BRUHIN elaborated. There are observations in support of this view, but no data demonstrating its validity.
Some authors hold the opinion that antlers are non-adaptive (LINKE,
I957), and that the animals would be better off without them (DARLING,
I937). Some have suggested that the antlers are now too large and complex
and have grown beyond the necessities of life (PORTMANN, I959; KRIEG,
from BENINDE).
A further thought upon horn evolution is expressed by HEDIGER (BRUHIN,
I953). He suggests that horns can be classified into three general functional
classes.
(I953)
I. Horns, similar in shape and size in both sexes function mainly as

weapons against enemies. e.g. Oryx, Hip potragus, and Bison.
2. Horns which are useless against predators and function only in intra-
I82
VALERIUSGEIST
specific fighting. e.g. Aepyceros, Cobus, Redunca. These forms have sexual
dimorphism, or horns in one sex only.
3. Horns which are never used as weapons, but have ceremonial function
only. e.g. Giraffa.
Let us turn to the most commonly stated function of horns, that is, as
weapons. This view is implicit in COLBERT'Sstatement cited earlier and
SIMPSON'S (I949) discussion on horn types of antelopes. There are several
dominant assumptions in these views. Not only that horns function as
weapons, but that they are similarly employed and have been guided in
their evolution by much the same forces. Their diversity is regarded as a
multiple solution to the same problem, which is to ward off enemies and to
dominate over conspecifics (SIMPSON, 1949). Horns are evidently conceived as stabbing weapons that evolved to inflict wounds on adversaries.
Clearly, had this been the evolutionary goal, then "one good pair of horns",
as COLBERTSOfittingly states, would have been all we could have expected.
It is interesting that both COLBERTand SIMPSON think that the country
in which bovids live is somehow relevant to horn evolution. This is understandable if one assumes that horns evolved primarily in response to
carnivores, so that horn bearers in one region evolved similar horns against
similar carnivores. Both COLBERTand SIMPSON noted however that great
diversity of horn types is found among antelopes inhabiting the same region.
The same observation can be made on Eurasian and North American
mountain bovids. No greater difference in horn size and shape could be
conceived than between the Caprinae and Rupicaprinae. Chamois and ibex,
rocky mountain goat and big horn sheep live frequently in the same habitat.
But not the slightest indication of convergence of horns can be detected.
For example, let us take Oreamnos americanus and Ovis dalli. Both are of
similar body size and may be found living side by side, although habitat
optima differ. The combined weight of the horns of large male mountain
goats is between Ioo and 120 grams. The horns from an old ram sheep weigh
about 6,900 grams, approximately 8 %oof his live weight.
The cervids offer rich material for arguments against the influence of
predators on horn evolution. Antlers are found in males only and are
functional for a very short part of the year. Male cervids lose their antlers
in winter at a time when they are harrassed most by predators. The study of
MECH (I963) showed that wolves harrass about I2 moose for every one they
kill. Without antlers the harrassed moose were capable opponents. Most deer
are well known for their ability to defend themselves with their forelegs,
rather than with antlers. Giraffes (BACKHAUS,I96I) also use their hind or
I83
front legs against enemies. For many small horn-bearing bovids like sheep or
goat, flight is a better protection from predators than fighting. There is no
evidence that wild sheep can fend off an attack by wolves, one of their
major predators. Furthermore, observations on Stone's sheep (Ovis dalli
stonei) and mountain goat (Oreamnos) revealed that these species are but
rarely found in regions of their habitat where a quick escape to precipitous
terrain is not possible. Only in those species which regularly meet the same
predator and do not flee, is there any hope of detecting an influence of
predators on horn evolution. The up-curving horn tip of musk oxen may
be an example. These bovids form a phalanx against sled dogs and presumably wolves, and are reported to hook at sled dogs with their horns
(PEDERSEN, I958). Species such as moose, though devoid of antlers during
most of the winter, may stand and fight harrassing wolves. In these the
rather terrifying defensive threat (GEIST, I963) may be a mechanism
primarily evolved as a predator deterrent. It is noteworthy that nothing
comparable to the defensive threat of moose has been found in any other
ruminant studied. Yet, this very threat type is common among carnivores.
There is little argument that horns can function as weapons and inflict
wounds on opponents. Yet this statement is a far cry from the generalization
that horns evolved as weapons. From the above discussion it appears unlikely
that horns evolved in response to predators, although in a very few cases
predators may have had some slight effect. Interspecific engagements appear
hence unlikely as an impetus to horn evolution. Intraspecific contacts therefore remain as the most probable functional influences.
THE EVOLUTION
OF SMALL HORNS
It is noticeable that horn-like organs appeared and evolved primarily

on larger mammals, less frequently on small forms. The horned rodents
(Mylagaulidae) are striking in their uniqueness. The tiny cephalopids and
South American odocoleinids carry small horns. These ruminants may
however be secondarily reduced in adaptation to the undergrowth they
inhabit. The duikers, for instance, have a surprisingly long gestation period
for their size (KRUMBIEGEL, 1954), an indication that they may have evolved
from larger forms. The most primitive ruminants, the small tragulids, have a
short gestation time and are quite hornless. In general, horns are confined
to animals of the stature of small deer or gazelles and larger. The circumstances for horn evolution are apparently less favourable in small than in
large mammals.
In large mammals a fighting form can become effective which would be
conceivably less potent in small forms. With increase in total size, a heavy
184
VALERIUS
GEIST
head swung in an arc or a frontal butt would deliver a substantial amount

of energy onto an opponent in animals the size of pigs, but very little in
animals the size of mice. The greater resistance of the large body to a heavy
blow would allow for a greater jarring effect, and for a greater chance
of tissue damage.
In early mammals of large size an evolutionary impetus would have been
present to adopt dealing out blows with the head as part of their repetoir
of fighting behaviour. Their size would have made large, violent body
movements difficult and dangerous to them. Rearing up would be precarious
to lumbering mammals, just as much as jumping at each other. A fall could
easily dislocate or break bones or tear muscles and tendons. Hence combat
forms, so easily and effectively used by small mammals and probably by
their earlier ancestors, would slowly but surely become an impossibility to the
ever enlarging beasts. Fighting with legs would be reduced by the necessity
of keeping all legs on the ground to support the heavy body. This leaves
mainly the head and neck free for action. The appearance of horns is
probably associated with a specialization towards dealing out blows with the
head, and concurrent reduction in other fighting forms.
When horns first appear in mammals, they are small protruberances on top
of the skull, located just anterior to the orbital region. We can see this in the
early titanotheres, dinoceratids and protoceratids. Their placement is such
that one way, and probably the only way, to use them successfully is to
swing the head up and sideways and deal the opponent a blow on the side.
These protruberances must have been adaptive, as they would concentrate
the energy of a blow onto a small area of the opponent's body. This would
be more painful, and hence more effective than a blow with an unhorned
head.
The descriptions so far are applicable to herbivores which show an increase
in bulk during their evolutionary history. Primary selective forces shape
teeth and legs in a manner which, respectively, allows efficient break up of
plant tissue and better body support and maneuverability. The tooth, jaw
and leg structures of carnivores are formed by different selection pressures.
These organs are not only efficient in dispatching prey, but also in maiming
rivals in combat. But the broadside intimidation display is common to
carnivores and herbivores alike and - as is made likely below- is an ancient
behaviour of mammals. If two primitive, large herbivores engage agonistically,
it would be while displaying the broadside to each other. We should visualize
them moving about each other, dealing out blows on the opponent's side,
belly and haunches. Their first line of defense being a tough, heavy hide.
There is good evidence for this hypothesis. The broadside engagement is
I85
very widely distributedin mammals.In some, however,it is found only as a

displayprior to engagement.Fights in which head blows on the lateralsides
are exchangedby opponentsare found in the Suidae (Sus scrofa) (MOHR,
I960; SNETHLAGE, I957), in giraffids (BACKHAUS, I96I), probably in
Okapiaas some elementsof this fighting form are describedin WALTHER,
I960, in bovids (GEIST, I964, in Oreamnos,rarely in Ovis dalli stonei).
Broadsidedisplays are a most commonoccurrencein agonisticinteractions
described in carnivores
(TEMBROCK, I962), bovids (WALTHER, I958;

I964), in giraffids (BACKHAUS, I96I; WALTHER,
I962b), in camelids(KOFORD, I957), in cervids (COWAN & GEIST, I96I),
and even in rodents (Rattus, EIBL-EIBESFELD, I963; and from personal
SCHLOETH, I96I;
GEIST,
observationson Citelluslateralis).The behaviourof many mammalshas not

been describedso far. Yet there is enough evidence to suggest that the
broadsidedisplayor engagementis a widely distributedor ancientbehaviour
patterncommonto the ancestorsof modernmammals.
The broadsideengagementin large mammalianconspecificscoupledwith
a headblowor a butt, conceivablygave impetus towards the evolution of
small horn-likeprotruberanceson top of the skull. The head blow can be
made more efficient not only by horns, but also by large, protrudingteeth
in the lower jaw. This course of evolution was grasped by the Suidae.
Hence, both the bony pedicle and the large tooth would fulfill similar
functions,and are analogousorgans in that context.
With the appearanceof horns or horn-analogousteeth, defensive mechanisms to reducethe efficiency of these organswere evolved.In present day
species known to employthe primitivelateralengagementwith head blows,
the first line of defence appearsto be a heavy, thick skin. Thus, the males
of Sus scrofa develop a very heavy connectivetissue shield on the lateral
sides of their bodies. The shield may be up to 6 cm thick (MOHR, I960).
During fights the opponentsblows are caught on this shield (SNETHLAGE,
I957). The shouldersand sides of wild male boars are frequentlyimpregnated with tree gum, which matts the long guardhairs and thick underwool,
forminga furtherprotectivelayer (SNETHLAGE, 1957). The AmericangoatantelopeOreamnospossesses a ratherthick, tough skin which was used by
Alaska Coastal Indians as a breast armour.The skin of Okapia is 5 to
6 mm thick in places and due to its toughnessvery reminiscentof thick
rubber sole (GIJZEN, I959). No published informationon the thickness
and consistencyof giraffe skin was found.
As horn-like organs and heavy hides evolved with the lateral fighting
engagement,several new roads opened to horn evolution. The thick hide
provided an evolutionaryimpetus towards sharper and harder horn-like
i86
VALERIUS GEIST
organs, to make the shield less effective. Equally, there is an impetus to

enlarge and strengthen the shield. It is probable that Sus moved along this
evolutionary road. However, damaging fights must be considered disadvantageous to both opponents. A wounded, though victorious male is no match
for equal sized opponents. If seriously wounded, he would soon drop from
reproduction due to illness as well as subdominant status. Sharp horn-like
organs are advantageous up to a point only, but then cease to be so. Display
and non-damaging fighting forms are selected since overt fighters do not
live long.
There are two ways to avoid overt fighting, or to reduce it to a minimum.
These are, to make an adaptive feature of an inhibition to strike conspecifics or, alternatively, to increase flight tendencies without restricting
agonistic action. Both of these ways are illustrated by members of the
Rupicaprinae, the former by the American goat-antelope (Oreamnos), the
latter by the European chamois (Rupicapra). Oreamnos males display
frequently but rarely strike one another. Powerful males accept damaging
blows from females without physical reply, and withdraw even from
agonistic juveniles (GEIST,I964). Rupicapra show a different mode of combat. Opponents alternatively rush head lowered at each other and avoid
each others' attacks by withdrawing. This leads to the aggressor chasing
the attacked. Suddenly the hunted turns and charges the aggressor who now
turns, flees and becomes the hunted. (WALTHER, i962; KNAUS I960;
ZEDWITZ 19I3), The chamoise buck is hence not inhibited from striking as is
the male mountaingoat. In both types of agonistic encounters, fights are
infrequent while displays predominate. The inverse relationship between
frequency of overt fighting and frequency of threat displays was first
suggested by WALTHER (196I) on the basis of his chamois observations.
Overt fighting can also be reduced by increasing minor agonistic engagements, which help to retain a stable dominance order. The maintenance
of such stability also requires that individuals in a troupe know each other,
that the troupe remains together, or that individuals inhabit limited home
ranges and know all other individuals frequenting that home range. It may
be that giraffes followed this evolutionary path. Giraffe bulls do engage
quite frequently in minor blow exchanges (BACKHAUS, I96I; INNIS, I958).
This is further substantiated by the heavily pneumated skull of giraffes
(LANKESTER, I902), an adaptation in animals that deal out heavy blows
with the head. The giraffe's horns are quite dull.
A further general means of reducing the effect of sharp, damaging
horns is to shift away from head blows or butts to an alternative manner of
fighting. WALTHER (1958) described the neck fight in the nilgau (Bosela-
EVOLUTION
OF HORN-LIKE
ORGANS
I87
phus), a short horned Indian antelope. Although the nilgau is fully capable
of butting, its intense threat display and neck fighting are probably a
means to reduce the primitive fighting form. It is noteworthy that nilgau
bulls face the opponent in a kneeling position while butting, and hereby
reduce the target area of the body which can be reached by the opponent's
horns.
It is postulated therefore that the first stages of horn evolution were
represented by small horns. These were associated with a broadside
intimidation display and with such defense mechanisms as dermal shields,
inhibition to strike, withdrawal or reduction of target area, early or quick
establishment of dominance order, or emphasis on non-damaging forms of
fighting in intraspecific engagements. These are perfectly functional solutions
as the presence of these forms today indicates. There is however no impetus
towards the evolution of larger horns. Large horns in association with
the suggested defense mechanisms have no conceivable advantage over
smaller horns. Hence the evolution towards larger horns is unlikely.
THE EVOLUTION
OF LARGE HORNS AND TUSKS
I have suggested a number of defense mechanisms against the blow of a

horned head. One extremely important mechanism was not mentioned. This
is to catch the opponent's blow with a counter blow and make it ineffective.
Regardless of whether the opponent butts or hits, if his head can be caught
and held, his attack fails. This defense mechanism requires a thick skull
and but a slight change of horn form. The early stages of this defense
form can be seen in Bos and Bison.
It is instructive to compare the agonistic behaviour of the American
goat-antelope Oreamnos, with that of Bos and Bison. Oreamnos represents
the primitive stage. The horns are short and pointed upward. The skull is
thin, very light and fragile. In contrast the skull of Bos is thick, with a
double roof of bone over the brain. There are large sinuses around the brain
case. In Oreamnos these sinuses are small and restricted to the region above
the cribiform plate. The horns of Bos swing out, then up. The intimidation
displays of Oreamnos, Bos, and Bison are very similar. All display laterally
to an opponent. Like Oreamnos, Bos and Bison turn the head from the
opponent while holding it low (FULLER,
I960; GEIST, I964; SCHLOETH,
in Oreamnos and Bison
arched
is
i96ia). The back
conspicuously
MCHUGH,
I958), and less so in Bos. If displaying while stationary, the
legs are closely bunched under Oreamnos and Bos. When approaching an
opponent, Oreamnos and Bison move at a slow, stiff walk and circle the
opponent. There are also differences. Bos and Bison tend to bellow,
i88
VALERIUS GEIST
whereas Oreamnos roars but rarely. Horning the ground and pawing or
wallowing are part of Bos and Bison agonistic behaviour. Oreamnos does
not wallow, while horning and pawing are infrequently seen in agonistic
interactions. In general, however, the actual displays of these three bovids
have many features in common.
During fighting, the target area for horn blows is identical in these three
genera. MCHUGH (I958) states that buffalos attempt to horn the flank of
the opponent. FULLER (1960) states that buffalo-bulls during fighting
attempt to deliver blows into the opponent's side SCHLOETH i96ia) writes
that in Bos, fighting partners are attempting not to lose frontal contact
until they can push the opponent into a position which allows them to
deliver a horn thrust into the body. McHUGH presents considerable evidence
showing that casualties from fights die of body wounds. These wounds are
located predominantly on the lower part of the body, and resulted from
horn punctures or tears. He recorded bulls with intestines protruding from
punctured bellies; several buffalos died from punctured and collapsed
lungs; some had punctured hind or front legs and were lame. On butchered
bulls McHUGH recorded that 23% of all bulls over four years old had one
to three fractured ribs. GEIST (I964) on the basis of field observations
on Oreamnos and the autopsy of two wounded males, showed that the
ventral body region is the target area for horn blows in fighting mountain
goat. As in buffalos, horn punctures and blows were located on the
abdomen, chest, as well as on the inside and outside of rear and front legs.
Oreamnos fights by swinging the head sideways; and the same type of horn
jab was observed in buffaloes by MCHUGH (1958).
Oreamnos, Bos and Bison hence show many similarities, yet their manner
of fighting differs conspicuously. This difference is clearly understandable, if the head to head engagements of Bos or Bisont serve as a
defense mechanism against attack on the vulnerable body. Whereas the head
evolved structures to absorb the shock of a frontal encounter, the outcurving
horns hold and control the opponent's head and prevent the flank attack.
In the evolution of horns and fighting behaviour, Bos and Bison should
be regarded but one step beyond the Oreamnos stage.
Not only the bovids have made a change from the lateral to the frontal
engagement. The Suidae have done the same. Sus is in many ways
comparable to the Oreamnos stage. Boars fight laterally, catching the
opponent's blow from the tusked snout on the dermal shield covering
shoulders and sides. As in Oreamnos, fights are damaging, as blood may be
found on the ground after some engagements. Boars shot during the rut
frequently show bleeding scars (SNETHLAGE, I957).
I89
The African giant forest hog (Hylochoerus) and the wart hog (Phacochoerus) are comparable to the Bos-Bison stage. Boars fight by running
head on into each other from some distance. Then follows head to head
pushing, while the opponents attempt to slash each other in the sides with
their lower tusks (MOHR, I960). If successful the lower tusks can cut
through the thick hide of Hylochoerus, and inflict deep wounds (D6NHOFF,
I942, from MOHR,I960). Hylochoerusand Phacochoeruspossess large upper
tusks which protrude laterally and curve up -very much like Bison horns
do. These tusks could not be primarily used for rooting - the popular
functional and evolutionary explanation for the wart hog tusks. The giant
forest hog, with tusks similar in shape to wart hog tusks apparently does
not root. Its nasal disk is covered by hair, which presumably would be worn
off in rooting. EWER (1958) was the first to recognize the large upper tusks
as organs analogous to ruminant horns. As in Bos or Bison, the curved upper
tusks of Hylochoerus and Phacochoerus help to hold and control the
opponent's head preventing the latter from using his sharp lower tusks
effectively. It may be that the spectacular tusks of Babirusa have a similar
function. In Phacochoerus the size of the lateral, dermal shield has apparently been reduced.
The analogy of horn-like organs should be extended to include the tusks
of elephants. KUHME (196I, I963) describes the engagements of captive
Loxodonta bulls. The animals move at each other with raised trunks and,
placing the bases of their trunks together, attempting to push each other
back. The tusks, if long enough, lie parallel along the opponent's head
(Fig. 2). If the weaker of the two opponents turns, it is regularly caught
in its side by the tusks of the stronger (KUHME, I96I). Hence, one of the
functions of the tusks must be to hold the opponent's head, and prevent latter
from bringing his tusks into action. Furthermore, tusks probably allow
effective pushing with the trunk bases, because they do not allow the
opponents' heads to slide past one another.
It is postulated that in the second phase of horn evolution, horns have
the function of a guard, as well as that of a bruising and piercing organ.
They help block an opponent's attack and make it ineffective. However,
a head to head contact of opponents easily leads to pushing and shoving.
This sets the evolutionary impetus toward the next stage of horn evolution.
THE EVOLUTION
WALTHER
OF COMPLEX HORNS
(1958, I962) on the basis of very extensive observations on
African antelope, proposed that the major function of horns was to guide
the opponents' heads together during the clash, then bind the heads to-
[90
VALERIUS GEIST
TPwo African elephants (Loxodonta) during an engagement. Note the placement

Fig.
of the tusks. They prevent any maj or sideways movement of the opponent's head, and
hence prevent latter from using the tusks effectively to damage his opponent.
gether in the following pushing and tugging match. The spirals, bumps and
ridges on the horns of various antelope and gazelles serve to hold the horns
together during the match, allowing the opponents to develop full strength
wrestling engagements. The shape of the antelope horns and mode of
fighting isi such that the horns are quite harmless to the conspecific
opponent. Although to human observers such horns may look immensely
dangerous (i.e. the oryx, WALTHER, I962).
Elaboration of horns from the simple Bos-Bison stage hence served
primarily to make pushing and wrestling matches more effective. There is
little danger to opponents from such fights, hence they may occur quite
frequently. This was noted for the impala (Aepyceros) by DASMANN&
MOSSMAN (I962),
who used elk (Cervus) and male deer (Odocoileus)
as standards for comparison.
I would suggest that WALTHER'Sideas about antelope horn function
apply equally to cervids. That is, cervid antlers should be regarded
primarily as structures binding opponents together during pushing and
wrestling matches. However, they must equally be regarded as guards
against the opponent's attack on his adversary's flank.
There is a surprising concensus of opinion among students of cervid
ENVOLUTION OF HORN-LIKE
ORGANS
I9I
biology, that males attempt to gore the body of their opponent. HENNIG
(I962) writes that roe-bucks (Capreolus) attempt to strike the opponent
ventrally, but also to catch the opponent's blow. RIECK (I956) writes that
roe-bucks, engaging from a lateral display, try to grip the opponent
ventrally with their backwardly directed antler points (see Fig. 1 m). He
cites an observation of a roe-buck slashing into an opponent's side when
it fell. KIDDIE (I962) in his extensive observations on Sika deer (Cervus
nippon) states (p. 23): "... the object of the fight appears to be to cripple
the opponent by goring him in the ribs or flanks." Sika stags fight fiercely;
the antlers take the weight of the charge. KIDDIE once shot one of two
fighting stags, whereupon the other gored his fallen opponent in the ribs
and stomach. KIDDIE also observed a fight between a Sika stag and a
small red-deer stag. He states: "The red stag could push the sika stag
where he wished, but was too slow to avoid the charge of the opponent,
which was attacking his ribs and flanks whenever he could. The red stag
seemed to be ushing and attempting to overthrow his opponent, while the
sika stag maneuvered and appeared to be waiting for a chance to deliver
a swift upthrust with the antlers. The sika stag bore severe wounds about
the neck and head, and the red stag was punctured in the stomach, with
some severe slashes on the shoulder that were bleeding badly. Now and then
the red stag attempted to run, but the sika stag was quick enough to thrust his
antlers at his opponent's unguarded flank, causing the red stag to turn and
fight once more." LINKE (1957) in his discussion of antler function in red
deer, also thought of them as organs binding opponents together during fights
and hence allowing a matching of strength, but he did not think this to be
their primary function. He also states that red deer attempts to gore
opponents into their side.
In deer, as in bovids, the body appears to be the primary target of
attack. In forms with elaborate horns however, the head itself becomes the
target in at least some engagements. WALTHER (I962) suggests that with
increased horn complexity went a greater tendency in bovids toward
frontal rather than flank attack. I concur with this view. However, I
suggest that in species with complex horns most attacks are caught by the
opponent with his horns (Fig. 3). Also, among social ruminants many
small engagements occur, which lack the characteristics of serious fights.
In such attacks the horns are indeed aimed at the opponent's head. Ovis
dalli stonei illustrates this well. During fights around the oestrus ewe, rams
forget their ceremonial head-on clashes. The defending dominant ram
attempts to crash into his opponent's body. Whirling around one another
rams in vicious fights hit out sideways and deliver blows on the opponents
Behaviour XXVII
I3
I92
VALERIUS GEIST
sides, shoulders, haunches, head, virtually any part of the body (Fig. 4).
These observations imply that during serious fights the body remains the
principal target area. The same is implied by BUECHNER'S (I963) observations on the Uganda kob (Adenota), (p. 6i); "'A third male may run in
to break up a fight by butting his horns into the side or hind quarters of
the combatants, sometimes inflicting severe wounds." It is apparent therefore that some bovids retain the orientation toward body attack while
directing most of their aggressive behaviour toward the head. Yet in some
species (i.e. Gazella tompsoni) only the head to head engagement may be
present (WALTHER, in print).
The above discussion suggests that cervid antlers and bovid horns evolved
convergently and function as weapons, secondly as guards and thirdly as
organs binding opponents together during pushing and wrestling contests.
Horns and antlers are elaborate. In the Bovidae however some groups
followed another road of horn evolution. We may term them the "rammers". The Bos-Bison stage serves as an equally convenient jump off point
for these as for the "wrestlers". Horns increased in size, though not complexity, and served not to bind the opponents together, but to deliver a
heavy blow. The sheep and goat, as well as the musk ox (Ovibos), are
excellent examples. But not only a heavy skull need be evolved to absorb
the shock of heavy bodies in head-on collision. MCHUGH(I958) points out
that the cushion of long hair on the head of Bison serves the same function.
He states that a bullet from a 30-o6 rifle fired from 30 feet into the head
of a bull, bounced back and was recovered a twisted piece of lead. In
principle the manner of battle in the "rammers" is the same. Conspecific
opponents clash head-on (Fig. 3). In particular fighting methods differ.
A conceivable evolutionary sequence begins with the Bos-Bison stage, where
opponents do butt heavily head on followed by some pushing. The next
stage is illustrated by Ammotragus or Ovis musimon. Opponents collide
head-on after a rush (BRUHIN, I953). The final stage has been reached by
Ovis canadensis, 0. dalli or 0. ammon poli (WALTHER, I96I), where
opponents rise and charge bipedally, then jump into the clash (Fig. 3).
THE EVOLUTION
OF HORNS AS DISPLAY
GUIDING ORGAN
OR ATTENTION
Observation of Artiodactyla reveal a trend toward cephalization of the

display organs (PORTMANN,1959). This proceeded for the most part
separate from horn evolution. Such display organs include striking hair
bunches, skin outgrowths and colour patterns. A comparison of the somewhat generalized Oreamnos with the highly developed head end of Bison
EVOLUTION
othersthey are cepalsdOrea
otes
hyar
epaied
OF HORN-LIKE
s()ilsrtedthe
remo
()ilutatdth
Fneig.
aneior
5.istnctsome
d.Istnction.gener
Iener ovi
ruiats,hespavy
Ofi ruiathe
ame-rworgns
dispavy
ORGANS
fome.'
ome.A
I93
oralhirrig
osl
ar
ig
are
dithiued
solerdisla
ditriued sover
thelaborgans
body,ans
I94
VALERIUS
GEIST
Fig. 6. A trend towards cephalisationof display organs is shown among some species of
Suidae. Sus scrofa shows a relatively undifferentiated head. SWuverrucosus displays not
only larger tusks but highly differentiated face warts. Phacochoerus shows the largest
tusks among living species of Suidae and is probablymore differentiated anteriorly than
any other suids (after MOHR, i960).
(Fig. 5) or of the plain head architecture of Sus scrofa in contrast to the

striking head specialization seen in Sus verrucosus or Phacochoerus (Fig. 6)
will illustrate the trends. It appears that in the Suidae there has been in
I95
parallel an increase in tusk size and in the large facial warts so prominent on
some members (Fig. 6). It can be anticipated that with a change in display
organs goes a change in display behaviour.
It was mentioned that many bovids, giraffids, some cervids and camelids,
whose behaviour has been studied, display laterally. In short horned bovids
like Oreamnos and Boselaphus, the whole of the animal's body is made
conspicuous during display. Let us look at the nilgau (Boselaphus), whose
display has been so meticulously described by WALTHER (I958). The bull
stretches head and neck stiffly forward and up. He appears extremely
tense. The steps are slow and stiff. The ears point back. The tail is depressed, but periodically is jerked up violently. Upon getting close the bull
assumes a full lateral stance to the opponent. To be noted are the colour
patterns on the bull's face, his dark mane, the conspicuous neck beard and
the white rings around his legs, supplemented by the tail flash. Any artist
would recognize on that bull the very principles of attention fixation and
guidance that he himself uses in painting. Small contrasting spots, sharply
projecting features, quick movements, localised jerks, quivers or flashesall these capture the attention of the viewer, and in the nilgau's case should
rotate the attention around the whole of the bull's body. The discovery of
edge receptors in vertebrate retinas (GRUSSER et al, I964), as well as
corticular cells in mammals that respond best to line or edge stimuli, or
moving edges (HUBEL, I963) supply the physiological basis for effectiveness
of attention-guiding adaptations.
If we look at the display in the Bison bull, we see him approaching his
opponent at a slow, stiff walk, while he may swing his head from side to side
(MCHUGH, I958; FULLER, I960). Due to the hair growth pattern on the
bull, most attention should be on the head and shoulder regions.
We recall the suggestion that horns functioned in display. Let us
contrast the display of small horned and large horned cervids. Little Capreolus
displays laterally to an opponent while raising his head very high (RIECK,
1956; HENNIG, I962). Odocoileus hemionus males display laterally to each
other while circling (COWAN & GEIST, I96I). From the lateral position
Capreolus and Odocoileus males suddenly clash and lock antlers. Both genera
are comparatively small-horned, excepting the largest of Odocoileus males,
and lack anterior differentiation as exemplified by Cervus (a mane), Rangifer (a mane) and A Ices (a skin outgrowth below the lower jaw, the bell). Nor
is there any report of the use of their antlers in display behaviour. However,
adult bull moose were on several occasions observed to stretch up the neck,
pull in the chin, tipping the antlers forward, thus displaying the light-colored,
huge palms, and to move slowly and stiffly at the opponent. This posture
I96
VALERIUS GEIST
was observed repeatedly by WALTHER (I963, personal communication) in

sika and red deer stags. During observations on rutting Rangif er bulls during
two years, it was repeatedly noted how caribou bulls circled tangentially
onto opponents, or cows that had wandered from the rutting band. It was also
observed that rutting bull moose approachedequal sized opponents tangentially
not frontally. Rangifer and Alces bulls would in this manner of approach
draw most of the opponent's attention towards their flashing antlers, and
white neck, or dangling bell. This circumstantial evidence hence suggests
that in some bovids and cervids, anterior differentiation was accompanied by
a behavioural change.
No consideration of display or display organs is meaningful unless they
are related in their function to the animal. We assume that they have been
developed through positive selective pressure and that they have evolutionary
significance. Displays are readily observable; the term implies intimidation.
The intimidatory function of display is often directly observable (Oreamnos
and Alces, GEIST, I964; Bosephalus, Oryx, Antelope, WALTHER, I958;
Bison, MCHUGH, 1958; Giraffa, BACKHAUS, I96I). But there is not always
a clear-cut predictable reaction to the display. The opponent may withdraw,
he may remain and seemingly pay no attention, he may attack, or assume
displaying himself. There is however little argument whether the lateral
display has intimidatory function, but rather if the hair ridges, outgrowths
and colour pattern contribute to the efficiency of the display.
Let us for the moment assume that display organs have intimidatory
function in conjunction with the display. It was postulated that cephalisation
of display organs had taken plate in some lines of ruminants and swine. It
is probable that scattered display organs are correlated with a broadside
display, and more cephalised display organs with a fronto-lateral display. If
this is so, then in species which possess anteriorly concentrated display organs
only, the broadside display should disappear altogether. This is precisely what
I have found in the North American sheep Ovis dalli stonei and Ovis canadensis. No trace of a functional broadside display was observed. Rather the
display is centered about the large horns of rams.
While dwelling in the company of companions, rams were frequently
observed to perform a peculiar behaviour pattern (Fig. 7). A ram would
move in a stretched posture through a band, head straight out. One could
observe one ram move to another in this manner. Sometimes equal sized
rams performed this pattern, called "'low stretch", to each other. Their
orientation was such that the open curl of the horn was pointing towards
the opponent. It was noticed when resting rams cocked their heads similarly
to the low stretching opponent, that again the curl was displayed. These
I97
observations suggested that the horns functioned also as display organs,

besides any other function they may have.
That ram horns evolved solely with the head-on collision type of fighting
isl not a tenable contention. Rams are the largest horned bovids for their size
and may carry up to io% of their body weight in horns. The horns grow a
sizable addition annually, but these additions decrease generally with the aging
of the ram. The longest horns therefore are in general carried by older rams,
and we find the ram population composed of individuals with various horn
sizes (Fig. 9). These graded horn sizes are not necessary for the fighting
pattern of sheep. Musk oxen also clash head on with terrific impact in their
fights, yet they grow a set of horns that does not change in size appreciably
with the age of the bulls. Bison may also clash head on, and we remember
McHUGH'S suggestion that the hair pad over the forehead functioned as
shock absorber. Unlike sheep, musk oxen and bison have a body display
(PEDERSEN, I958). These were reasons enough to investigate whether ram
horns function as display organs.
If the horns of rams were indeed selected for size and functioned as
display organs, then several conditions must be satisfied:
I. It must be shown that the low stretch is comparable to the displays of
other ruminants. The low stretch must be predominantly shown by the
more dominant male. It should have the same effect as the broadside display.
2. Rams must be aware of horn size differences of conspecifics.
3. Individuals with the largest horns must enjoy some advantages.
4. Large horned individuals must reproduce more than or to the exclusion
of small horned individuals.
5. Large horns must carry dominance status irrespective of the individual's
age.
6. The large horns, once lost, lead to a reduction in dominance rank.
The data following was gathered on Stone's and Bighorn sheep between
I96I and I964. Quantitative data will be presented only for the Stone's
sheep gathered in I96I-I963 in northern British Columbia. Only points 2,
3 and 4 will be illustrated with quantitative data. Data applicable to other
points will be published later.
To point I.
I have found that the low stretch is shown predominantly by the larger to
the smaller ram (Fig. II). There were no clearcut actions of the opponents
indicating an immediate intimidatory function of the low stretch. However,
smaller rams would react in the fashion of subdominants.
198
VALERIUS
GEIST
To point 2.
Rams were classified on the basis of horn size into four categories. Rams
whose horns formed a complete curl or more made up Class IV; rams
with horns less than full curl but with at least 94 curl heads were put into
Class III; rams with less than /4 curl but more than '4 curl made up Class
II; rams with '4 curl or less were placed into Class I (Fig. io). Classes
arranged in order I to IV showed a progressive increase in horn size and age.
An increase in body size was noticeable up to Class, III; a change in facial
features due to increase in thickness of horn bases, up to Class III. Class I
represented rams 17 to 25 months of age, Class II 2Y2 to 5 year old, Class
III 5 to 9 years old, Class IV 7 to 14 years old. Usually rams reached the full
curl stage at age 8 to 9 on the study area. The greatest change from Class I
to IV is shown by the horns. These data refer to Ovis dalli stonei.
ioo%
7.5
25
+3
Fig.
ii.
t2
+1
-1
-2
Ordinateequalspercentfrequencyof sexual approaches.

Abscissaequalssize
gradients of rams. Here 0 means same size class; about 70% of all approachesto the
samesize classweresexualapproaches.
+ i and- i meanthe next largersize class,and
the next smallersize class respectively.

It canbeseenthatlargerramswerelessfrequently
approached
sexually,and smallerramsmore frequently.This indicatesthat rams could
differentiatethe some what arbitrarilyestablishedhorn size classes.
Whether rams can differentiate each other on the basis of horn size, can be
answered by at least three different methods. The first is the multiple choice
method. Does one class, of rams prefer any other class to interact with and
choose that class more than any other class? Secondly, preference of one
class by the other is revealed by the frequency of patterns performed by one
class on or towards another. Thirdly, rams show a priori before an engagement whether they will act as dominant or subdominant individuals. In
7,~~~~~~~~~~~~~~~~~~,A
U1.~~~~~~~~~~
aYaS~~~~~~~~~~~~~~~~~~~~
Fig. 3. The main fighting form of the "rammers"is the head on clash, here i
by Ovis caniadensiscatiadensis. This photograph illustrates several other poi
attack was initiated by the smaller ram on the left. The larger is seen in a turn j
to catching the opponent'sblow with his horns. The attack was hence aimed at
ram's body, not head. It is also evident that Ovis canadensis jumps into the cl
hindlegs of the ram on the left are in mid air. (All pictures of sheep taken in M
in December) showing rams in the initial stages of moult).
It
Fig. 4. During vicious fight (occurring mainly around the estrous

around one another dealing out blows on any part of the opponen
landedon the opponent'sshoulder.It can be seen how both rams turn
attempting to put the heavy horns between the opponent and thei
heavy horns serve to catch blows, as well as for dealing
NE~~~~~~~~~~~~~~~~~~~~~~~~~~~~~7
'~~~~~~~~~~~
~ ~
.
Fig. 7. The low stretch or horn display of sheep, here illustrated

canadensis. Characteristically, the action of large, domi
:, .:
t
'*
ij(llC(:
i ,:::
W^:
re..
su
::: P
jr ci:I
*:
-+
**.- .X i^ M
,- ;:s
?
...
.,00,.
..
- VJ
'-*
j "."
***'.
.-*.
'j
,L*
,,
-IldCC
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LY
**
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'j*:?
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*;.r*i.
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3
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SkCL-
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*rr,
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:"C
*
Fig. 8. The horn threat is shown typically by a smaller towards

former initiates an engagement. The 3 year old ram on the left in
the larger ram lowers his head to meet the appr
Si,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
Fig. 9. Ram bands are assemblies of individuals, three years of ag

variable horn sizes. Two Glass IV rams (full curls) are seen at le
Class II rams.
.
_--J.
- ''t
~~~~~~~~~~~A
?N?.?
Fig. io. (a) A two,year old ram, Class I, (b) a three year old Class II ram, (c)
7 year old Class III ram, (d) a Class IV ram, well over io years of age. Thes
are of correct relative size. Note, although body size increases up to Class III a
the rna'or-change in the appa-nt characteristicsof rams is brought about by horn
EVOLUTION
O1
HORN-LIKE
ORGANS
I99
my studies all three approaches have led to the same result but only the
third method will be discussed here in detail.
Rams do not differentiate in their behaviour between males and females.
Rather, they differentiate between sheep larger or smaller than they are.
All smaller sheep, irrespective of their sex are treated like females, whereas
all larger sheep are confronted with agonistic behaviour. For instance, a
large full curled ram will treat a sub-dominant full curl like an ewe, while the
latter reacts with agonistic patterns. The behaviour performed towards a
smaller sheep will be termed "sexual", although for present purposes this
term serves as no more than a nametag! Preceding an interaction,
a ram already shows during the approach whether it has recognized the size
of its opponent. Towards smaller sheep the initiator adopts usually the low
stretch (Fig. 7). This is frequently followed by a sexual pattern. Towards
larger rams the initiator usually shows a horn-threat, followed by an agonistic
pattern (Fig. 8). Neither sexual nor agonistic treatment will be discussed
here further but will be treated in detail in a later paper. The important
point is the following: Using the first two behaviour patterns shown by a
ram initiating an engagement, we can classify his approach as sexual or
aggressive.
We derive the percent frequencies of sexual approaches shown by each
class towards all others. Next we add the percent frequencies for the following situations: all classes approaching the same class; all classes
approaching the next smallest class, the second smallest, the third smallest
class; all classes approaching the next largest class, the second larger and
third larger class. The frequencies are averaged for each situation. The result
is shown in Fig. ii. Next we graph the difference between the sexual
approach frequencies for one situation and the next following. This is shown
in Fig. 12. The interpretation of this graph is that a ram is most certain of
his actions towards larger or smaller rams, but least certain towards rams of
his own size. Horn size is the major difference between classes and it seems
valid to conclude that rams are indeed differentiating horn size classes.
To point 3.
It must be shown that rams enjoy advantages solely due to their horn size.
Table I shows that smaller horned rams have a greater tendency to withdraw
from larger horned rams. However, these data would not fulfill our requirements. It could be argued that the superior fighting ability of larger rams
accounted for the more frequent withdrawals of the smaller. After all, rams
in constant association with each other do set up a dominance order. Table I
may show no more than the result of the dominance order. Clearly, if horn
VALERIUS
200
GEIST
>~50-
S,53020 -
10
I
gXI
+2
+3
+1
-1
-2
-3
2
larger
sizeclasses
smaller
Fig. I2. Abscissa and ordinate same as Fig. i i. Here the difference in the percent
frequency of sexual approaches is graphed. The graph is interpreted as showing that
rams are more uncertain of how to behave towards equal size opponents than towards
smaller or larger rams.
lol
s40-
-e
X30'
z
H
20-
10-
+ 3
+ 2
+ 1
larger size classes

2
-'1
-2
-3
smaller size classes

2
Fig. I3. Abscissa as in Fig. ii and 12. Ordinate shows the numberof behaviourpatterns
of sexual (black) and agonistic (white) nature receivedby a strange ram entering a band.
It can be seen that he receives most patterns from rams of the same size class, and least
by much larger or much smaller rams.
size carries some advantage, it must show itself before a dominance order
based on actual physical interactions is established. The large horns of full
or three quarter curled rams should derive for their bearer a priori some
TABLE
20I
Withdrawaltendenciesof rams (Ovis dalli stonei; I962 data)

Interacting One ram is One ram is
ramsof same one horn
two horn
horn size
size smaller sizes smaller
Number of interactions
I36
73
134
No. of interactionsin which one ram

turned from the opponent -
or
movedaway
No. of withdrawalsin which a ram
movedaway, ratherthan turnedaway
only from his opponent.
30 (22%)
(I3%)
37 (28%)
2I
(57%)
3I (43%)
30 (97%)
degree of dominance, in an association of strange individuals. Ram bands

are not stable associations throughout the year, but rather they fragment and
reassemble from different individuals at specific periods!of the year. Hence
during the annual cycle a ram periodically meets a group of strange rams
and must establish his dominance position. If his horn size does give him some
measure of dominance, then he should be found fighting only with rams of
the same horn size class. Other rams! should, during the first meeting,
recognize his attributes by his horn size, not having as yet experienced any
body contact with him. The desired information can be only attained from
known rams in a group all members of which are known to the observer.
Then, one must be fortunate enough to catch a ram just after a major
movement, when it begins interacting with strange rams. Such observations
are by their very nature rare and hard to come by. I have been able to
observe the integration of only 7 rams into groups of new individuals and
could gather quantitative data for five. In every single instance these rams
interacted extensively only with rams of the same horn size class, and were
largely ignored by rams of different horn size classes. This is demonstrated
in Fig. I3.
The quantitative data tells but a fraction of the story. In fact, it appears
from the graphs that some rams have not entirely recognised the attributes
of the new companion. For instance, recorded in the data is the following:
A strange (full curl) Class IV ram appeared in one of my ram bands under
observation. A Class III ram approached and displayed dominantly (using
sexual or courtship patterns!) to the full curl. This lasted but a few moments,
for the full curl whirled and rose into the bipedal threat display. The Class
III ram ran away. The dominance order was settled without a fight. Yet
VALERIUS
202
GEIST
on what basis had the Class III judged his opponent? Both were 7 years old,
and of very nearly the same body size.
Not included in the data are observations on one very large full curl and
a small 4 year old Class II ram that crossed to one of my observation bands.
In the following seven days but one single engagement was observed between
that full curl and any member of the band. That engagement was with a
large Class III ram who acted a priori agonistically rather than sexually,
indicating that he had submitted. None of the large rams from the observation
band ever approached this large stranger. The little 4 year old Class II ram
remained close to the large full curl for several days. The very day he was
observed without his large companion, two 5 year old Class II rams had
engaged him. This engagement was partially filmed.
It must next be shown that large-horned rams are more likely to reproduce
than smaller horned rams.
3.0-
(a) L Mountswhichledto copulation

(b)
All mounts
.0
2.0-
1.0
IV
d
I
III
Hornsizeclasses
Fig. I4. Ordinateequalsmountsperformedby ramson oestrousewes per hourspentin

the vicinity of oestrous ewes. Abscissa equals horn size classes of rams. It can be seen
that most mounts were performed by the large horned full curls, and least by the small
horned yearling rams. As horn size increases, so does frequency of mounts on oestrous
ewes.
To point 4.
During the sheep rut of I96I and I962, Stone's rams were observed for
46 ram-hours in competition around estrous ewes. The distribution of
EVOLUTION
OF HORN-LIKE
ORGANS
203
ft'W
Fig. I5. The evolution of horns in bovids. The primitive horns are short, sharp. The
skull is thin and fragile; engagements are lateral and broadside display prominent
(Oreamnos, GEIST, I964). Next the head to head combat evolved as a defense mechanism.
The skull is heavy, horns curve out to hold opponent.Broadside display prominent (Bos,
SCHLOETH,I96I). From the primitive frontal engagement two combat forms evolved,
the "wrestlers" illustrated by Antelope (WALTHER,I96Ib). Combat form a frontal pushing
match allowing test of opponent'sbody strength. Broadsidedisplay prominent.The second

combat form
is illustrated
The 09:57:34
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and function.
Conditions The broadside display
204
VALERIUS GEIST
mounting frequencies for various ram-classes are shown in Fig. I4 (a) and
(b). (b) is a record of all mountings, be they interfered with or not. In (b)
mountings are included which could not have led to copulations due to two
factors, one the withdrawal of the ewe, two the interference by the old, full
curled rams. The data shows clearly that the full curls perform most of the
copulations.
Not included in the graphs, but very apparent in the field, is the observation that small rams are able to chase away and steal the ewe only after the
guarding, large ram has copulated frequently and appears exhausted. Or,
the large ram has stopped guarding, feeds and rests, while the next largest
ram commences guarding and copulating with the ewe. This is usually a large
Class III ram.
To point 5.
If large horn size carries evolutionary significance, large horns must
carry dominance status irrespective of the individual's age. It is very readily
apparent in bands of rams that among the full curls for instance there may
be a sizeable number of younger rams. In the study area for Stone's sheep the
largest and most dominant rams were in fact all 7 and 8 year old full curls.
Hence dominance is not entirely age dependent. We saw that horn size
correlated closest with dominance rank. Hence, horns carry dominance
status irrespective of the individual's age.
To point 6.
If the large-horned dominant loses his horns, he suffers a reduction in
dominance also. This proposition could not be investigated. Sheep carry
large, permanent horns and unlike deer do not lose them. In cervids the
above proposition could readily be investigated. Observations by BRUHIN
(I953) on fallow deer (Dama) show that the loss of horns can lead to loss
of dominance. A similar observation was made on moose (GEIST, I963).
COLLIAS (1956) observed that in his herd of goat (Capra hircus), individuals
with horns regularly dominated individuals without horns. Loss of horns in
the Oryx need not lead to a loss in dominance (WALTHER, I958). There is
however, no evidence, direct or circumstantial, that oryx horns have display
function. On the contrary, the indirect evidence points to these horns as
binding organs only.
On the basis of the reported date I conclude that sheep horns assumed
among other functions, the function of display organs. Hence horns have
evolved from a function of weapons only, to assume, in various genera, the
additional functions of guard, binding organ, or display organs (Fig. i6).
EVOLUTION
OF HORN-LIKE
ORGANS
205
HORNLIKE ORGANS AS RANK INDICATORS AND THEIR

RELATION TO TERRITORIALITY
We do not find graded horn sizes among adult males in many species of
ruminants. This condition is found among the large cervids, and among
the caprids. But by far most species of ruminants feature adult males that
differ little from each other (Fig. I5). The question arises, why did some
species evolve with horns that change conspicuously with age, while others
did not? In sheep it was, shown that graded horn sizes led to predictable
social relationships and a reduction of overt fighting. This must lead to
substantial savings in energy. This could be highly adaptive for large cervids,
k.
~ ~
Fig. i6. A male of the Uganda kob (Adenota). In this species no conspicuoushorn size
gradients as in Ovis exists. Males of the Uganda kob are territorial during mating and
show a lech type of breeding (see BUECHNER I96I; I63).
for instance, living in temperate climates. Males live off stored body energy
during the rut. The less energy expended during the rut, the more is left
for the strenuous winter following. I postulate that in species with graded
horn sizes males learn to associate large horns with superior strength during
frequent, minor agonistic engagements outside the rutting season. Males
fight extensively only with equal-size opponents, hence saving energy by not
206
VALERIUS GEIST
fighting all males. The saved energy can be used during the following winter.
For sheep there is substantial evidence that increased energy expenditure
during the rut is associated with increased winter-mortality (GEIST, unpublished). Frequent minor agonistic interactions between males prior to the
rut were reported by SCHLOETH (i96ia) for red-deer, and GEIST (I963)
for moose. Stone's or bighorn rams engage daily in minor interactions. The
hypothesis outlined above indicates the evolutionary "raison d'etre" for
graded horn sizes, but it does not explain why other species feature adult
males that differ little from each other.
If not asocial, agonistic animals live in dominance hierarchies. These
establish priority of actions and possession. They allow individuals to live in
a predictable social environment, as each can anticipate specific actions from
dominants and subdominants. They allow increased exercise and selection
for memory. Dominance hierarchies allow stability in groups of animals,
which ensures the least energy expenditure towards social needs, and a
maximum of energy devoted to growth, reproduction and individual wellbeing.
Dominance hierarchies can exist in groups of individuals that differ
little from each other. They depend on the knowledge of each individual
about the agonistic potentials of his companions. The society of most
primates appears to be of this type. Among ruminants cattle may dwell
in such societies. Both SCHLOETH (I96I) and MCHUGH (I958) report groups
of individuals uniting against one companion - a good indication of closed
societies. These individuals live in very predictable social environments as
long as the troupe remains together and recruitment is from the inside. There
is however, no predictability of social relationships if a number of strange,
similar individuals are united. Such predictability is preserved however, if
some sign system exists which parallels fighting potential, and hence dominance order. In sheep (and probably large cervids) horn size parallels
dominance order. Hence predictability of social relationships is preserved
even if strange individuals meet. In species with a rank system based on
horn or body size one can expect open societies, in which troupes periodically
assemble from strangers. This is what happens to bands of Stone's sheep.
However, rank systems can be based on criteria other than horn or body size.
Recent studies on African antelope and gazelles established the existence
of territoriality in Alcelaphus (BACKHAUS, I958), Adenota (BUECHNER,
I96I), Aepyceros (DASMANN & MOSSMANN, I962), Gazella thomsoni
(WALTHER, in print), and temporary or periodically shifting territories in
Gorgon (TALBOT & TALBOT, I963). In all these species adult males are
very similar in appearance. The same is true of the territorial South
EVOLUTIONOF HORN-LIKE ORGANS
207
American camelid, the vicunia (KOFORD, I957). In all these social, territorial artiodactyls, territory holders make up but part of the total male
population, while non-territorial males form associations on communal
areas.
Territory-guarding males are very conspicuous on their territorial grounds.
Gorgon bulls assume a very erect head posture and a rocking gait while
circling the female herds. Non-territorial bulls do not assume these striking
behaviour patterns (TALBOT & TALBOT, I963). Adenota males show
ritualised display towards approaching rivals (BUECHNER,I963). Gazella
thomsoni marks its territorial borders extensively with glandular secretion
(WALTHER,in print). The status of territory ownership is hence signalled
to conspecifics. But territory ownership is associated with a high dominance
rank. Hence, in these species, where males are similar in appearance, territory, rather than horn or body size signals a predictable agonistic
potential.
The extensive olfactory marking observable in territorial forms should
be thought of as an olfactory intimidation mechanism, as was independently
suggested by WALTHER(I96I), and GEIST (I964). This mechanism would
still be functioning in the absence of the owner. Olfactory, visual and
auditory signs produce to the conspecific an unmistakable situation, which of
course, allows predictable social relationships. Without this, an animal would
not live in a predictable environment, which after all, its adaptations
anticipate. It is suggested that territoriality in ruminants evolved as a
mechanism ensuring predictability of social relationships. Territoriality and
horn size gradients should be thought of as compensating. Both are rank
systems which parallel fighting potential and hence dominance order. Sexual
as well as age dimorphisms may equally function as rank systems. One of
the main significances of rank indicators is probably that they allow the
existence of "open"societies, where strange conspecifics can integrate with
a minimum of combat, damage and energy loss.
Yet why should there be displays and display organs, dominance hierarchies and rank systems? What evolutionary reason perpetuates these
universal phenomena?
The mechanisms in question protect and retain two utterly essential
attributes of life - self assertion and aggression. By the former I mean the
individual's energetic and egoistic attempts to satisfy its appetite relating
to food, reproduction and habitat selection. Aggression arises from self
assertion when one individual attempts to displace another during conflict
of mutual interests. Aggression is, adaptive. It ensures the more aggressive
having priority in satisfying its basic appetite; dominance hierarchies
Behaviour XXVII
I4
208
VALERIUS
GEIST
establish such priorities in groups of individuals. An individual without self

assertion is not only non-adaptive but unthinkable. Yet, overt aggression,
fighting, is for most mammals highly unadaptive.
The luxury of fighting is but possible to species with excellent defensive
mechanisms or ineffective weapons which allow only bloodless combat. To
others fighting is dangerous. A very aggressive individual is likely to provoke
a desperate defense once he damages a conspecific by engaging bodily.
This virtually insures that both combatants are damaged. Though the
aggressor may dominate, after a fight or two, he is no match to an uninjured opponent; his wounds may become infected leading to sickness, loss
of dominance, reduced reproduction and possibly shorter life span. Adaptive
though aggression can be, in its extreme form it is not.
Display or threat are mechanisms achieving the same end as overt
aggression, but not the same consequences.
I postulate that displays have their origin in stimulus contrast which
triggers escape in conspecifics. Selection greatly favours individuals which
provoke a major, sudden change in the vicinity of the opponent. Such
major environmental changes, or stimulus contrasts, are sudden fast
movements, loud, harsh and discordant sounds, and penetrating pungent
odours. I am assuming here that withdrawal is innate towards any major
environmental change in living beings; such stimuli signal basic dangers
of abiotic (i.e. rolling rock or falling branches) and biotic nature (i.e. predators). Two further groups of signals are postulated as adaptive. The
individual performs the initial motions of combat before his opponent,
and brings his weapons into position for use. This allows the opponent to
predict impending events. It requires though, that the opponent possess
knowledge of combat. Secondly, larger than actual size can be faked by the
aggressor. Withdrawal of smaller from larger is virtually universal, and
said behaviour would be selected for. The universals of strong but not overt
aggression are hence stimulus contrasts, intent to use weapons, and
increase in apparent size.
It was previously mentioned how defensive mechanisms evolved against
short but sharp hornlike organs. One set of selection forces favoured
efficiency of the weapons, while others favoured the efficiency of the
defenses. Hence two antagonistic sets of selection pressures were at work,
one nullifying the gains of the other. Similarly, in intimidation displays
there would be forces making the displays more effective by: increasing
apparent or real height, selecting for attention guiding adaptations, and
making offensive organs more conspicuous, while others would select for
mechanisms of psychological or behavioural nature, making these displays
209
less effective. If A displays, it is not advantageous for B to withdraw at

once, but only when a fight is imminent. By resisting and counterdisplaying, B may, in fact, move A to withdraw, and hence gain the fruits
of aggression. Such selection forces would lead to intense display performances in species that fight damagingly. This is precisely what is found in
Oreamnos, Rupicapra or Ursus (D. KLEIN, personal communication).
Though it is unthinkable how forces could act making a primary adaptation
like a tooth effective while others work to make it ineffective, for social
adaptations agonistic selection forces are logical.
Furthermore, I postulate that a conspicuous, though not major, difference
in an individual's appearance or behaviour, is likely to make that individual
more intimidating in pre-combat interactions, than normal ones. Such a
difference, since adaptive, would spread through a population. The effect
of this difference would diminish as its frequency in the population
increased, due to habituation of individual. Hence a population of uniform
(or nearly so) individuals with said difference is at the same stage as a
uniform population without said difference. Hence the potential constantly
exists that a conspicuous difference in appearance or behaviour spreads
through a population. The above is an unverified set of premises as yet.
However, if they are true, then (a) intimidation mechanisms have evolved
for all sensory pathways, (b) morphological social adaptations are changing
steadily and more rapidly than primary adaptations evolved for habitat
exploitation. In short, while the animal remains basically the same it will
change continually in outward appearance. It follows that two populations
arising from one, while basically similar, will differ increasingly in
appearance with the course of time. Small changes in outward appearance
will accumulate and because these changes are unlikely to be the same in both
populations, their appearance will differ. The evidence for points (a) and
(b) will be examined in a later paper. The present aim is but to present a
plausible hypothesis for the evolution of displays and adaptations making
these effective (display organs). We must attempt to understand these
universal, conspicuous phenomena, of which hornlike organ form a part.
A major weakness of this paper is that it does not explain adequately
the diversity of functionally similar horntypes,. I purpously refrain from
a detailed discussion of this topic. As yet we know the fighting behaviour
of relatively few ruminants and have really detailed descriptions on fewer
still. It cannot be decided at present that the deviations of hornforms are
not somehow related to small variations in a similar mode of combat.
One can explain the diversity of hornforms by postulating that hornshape
is not sharply selected for, rather it is modified by the pliotrophic effects of
210
\TALERIUS
GEIST
the species genes. As the genome of the species changes so does its hornshape since different genes would indirectly influence the horngrowth
characteristics differently. Individuals of any one species would carry similar
hornshapes, since they would have similar gene compliments. This view
explains hornshape, but it is all but untestable.
HUXLEY'S(193I) view of heterogenic growth serves to explain relative
hornsize differences between small and large forms. Unfortunately, this
explanation is faced by many exceptions. The interested may for instance
compare the horn and bodysizes among the various genera of the closely
related Caprinae and Rupicaprinae or ponder the hornsize differences between
Cervus unicolor and Cervus canadensis, or between Bibos and Bubalus,
or between Odocoileus and Hippocamelus. If in a species evolutionary
history hornsize increased with bodysize according to the laws of heterogonic growth, it appears to me more likely due to the adaptive value of
large horns than to the accident of heterogonic growth.
SUMMARY
Hornlike organs evolved independentlyin a number of mammalian families. Though
these organs assumed great diversity they did evolve into several general functional
types. A short review of the structure and development of hornlike organs is given.
Some views on horn function and evolution are critically discussed.
The evolution of hornlike organs is visualised as follows:
In primitive large mammals the head blow became effective as a fighting form due
to increased mass and inertia of the heads. Some forms grasped this potential. Combats were carried out from the broadside while opponents delivered head blows on
each others body. Skull protuberancesnow became adaptive. Concurrently, defensive
mechanisms evolve, decreasing the effectiveness of these protuberances. Foremost
among them is a thick, heavy hide or specialised dermal shield. These adaptive syndromes gave no impetus towards larger and more complex horns.
This impetus arose with the appearanceof a new method of defense -catching the
opponent's blows with the horned head. This leads to the evolution of heavy skulls
and horns capable of catching and holding the opponent's head. The target area of
attack remained the body. Frontal engagements resulted from the opponents' attempts
to control each others horned head.
It is shown that bovids and suids followed similar evolutionary roads in their mode
of combat.The tusks of the suidae fulfill the same functionand were subjectto
similar selection as the horns of short horned bovids. Thus Sus and Oreamnos,and
Bos and Phacochoerusare entirely similar in their mode of combat, hornlike organs
and defense mechanisms.
The primitive frontal engagement gave rise to two different modes of combat,
ramming and wrestling. The "wrestlers" evolved complex horns, whose function is to
bind opponentstogether,thus allowingthem to developfull strengthwrestlingand

pushing matches. The "rammers"evolved a heavy skull and heavy horns.
A trend towards cephalisation of display organs can be detected in ruminants and
suids. Display organs are interpreted as attention guiding adaptations. It is shown
that the distributionof displayorgans correlateswith the display.A broadsidebody

display is found in ruminants with a diffuse distributionof display organs. lf display
organs are cephalised, the broadside display disappears. This is found in the North
EVOLUTION
OF HORN-LIKE
2II
ORGANS
American wild sheep. Display centers about one heavy, very large horns of males.
Next it is verified that the rams horns are display organs. It is shown, the horn
display is the prerogative of the dominant; sheep can differentiate horn size classes;
large horned males enjoy reproductiveadvantage; horn size carries a priori dominance
status.
Hence horns evolved to function as weapons inflicting damage; as defense organs
shielding their owner; as binding organs allowing opponents a secure lock in battle;
as display organs having an a priori intimidating effect on certain conspecifics.
Some species of ruminants possess hornlike organs which change progressively in
size with the age of their owner. It is postulated that, as in North American wild
sheep, horn size parallels dominance order. Conspecifics learn to associate lhorn size
and agonistic potential of opponents. Hence horn size signals to conspecifics predictable
social relationships. In large cervids from temperate and cold climates this system
must save stored energy during the rut, which males need for the severe winter.
Graded horn sizes allow wild sheep to live in an "open" society in which strange
individualsmay meet and fit into the dominancehierarchy with a minimum of combat.
Graded horn sizes hence serve as rank indicators. It is next made plausible that
territories in ruminantsfunction in conjunctionwith displaying males as rank indicators.
Outside the territories males mix freely. Hence graded horn sizes and territoriality may
enable "open" societies, by acting as communicationmechanisms and creating a predictable, social environment.
A hypothesis is presented explaining the selective forces for the evolution of display.
It predicts that, antagonistic selection forces shape display and display-organs, that a
potential exists for rapid spread in a population of differences in behaviour and
morphology, that intimidation mechanisms evolved for all sensory pathways, that external appearanceaf animals changes more repidly than their primary adaptations.
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MEISTER, W. (I956). Changes in histological structure of the bone of white-tailed deer
during growth. - Anat. Records ii6, P. 709-72I.
MOHR, E. (i96o). Wilde Schweine. - Die Neue Brehm Biucherei.A. Ziemsen. Wittenberg-Lutherstadt.
PEDERSEN, A. (I958).
Der Moschusochs. - Die Neue Brehm Biicherei. A. Ziemsen
Wittenberg-Lutherstadt.
PORTMANN, A. (I959). Einfiihrung in die Vergleichende Morphologie der Wirbeltiere.
- Basel.
RIECK, W. (1956). Das Rehwild. - In: F. von Raesfeld. P. Parey. Berlin.
RYDER,M. L. (I962). Rhynoceros Horn. - Turtox News 40, p. 274-277.
SCHLOETH,R. (i96ia). Das Sozialleben des Camargue-Rindes.Z. Tierpsych. 12,
P. 574-627.
EIBL-EIBESFELD,
EVOLUTION
OF HORN-LIKE
ORGANS
2I3
(1i96ib). Einige Verhaltensweisen im Hirschrudel. - Rev. Suisse. Zool. ig6i,

P. 243-247.
SIMPSON,
G. G. (I949). The meaning of evolution. - Mentor Book. New American
Library. New York.
SNETHLAGE,
K. (I957). Das Schwarzwild. - P. Parey. Berlin.
SODY,H. J. V. (I959). Das Javanische Nashorn. - Z. Saugetiere 24, p. I07-24I.
TALBOT, L. M. & TALBOT,
M. H. (I963). The Wildebeest in westem Masailand, East
Africa. - Wildlife Monographs.No. I2.
TEMBROCK, G. (I962). Versuch einer Analyse des Imponierverhaltensbeim Rotfuchs,
Vulpes vulpes (L). - Z. Tierpsych. I9, P. 577-585.
THEMUS, E. & HOFER, H. (i96o). Stammesgeschichte der Siugetiere. Springer
Verlag, Berlin.
WALTHER, F. (I958). Zum Kampf- und Paarungsverhalten einiger Antilopen. Z.
Tierpsych. I5, P. 340-380.
(ig6oa). ,,Antilopenhafte" Verhalfensweisen im Paarungszeremoniell des Okapi
(Okapia johnstoni Sclater, I9oI). - Z. Tierpsych. I7, P. I88-2IO.
(ig6ob). Beobachtungenzum Sozialverhalten der Sasin. - G. v. Opel Freigehege
fur Tierforschung JahrbuchI959/I96o,p. 64-78.
(I96I). Einige Verhaltensbeobachtungen
am Bergwild des Georg von Opel-Freigeheges. - G. von Opel Freigehege fur Tierforschung Jahrbuch i96q/6i, p. 53-89.
(i962a). Von Waffen und Kamnpfenbei Horntieren. - Die Blauen Hefte 3/4.
(i962b). tber ein Spiel bei Okapia johnstoni. - Z. Siugetierkunde 27, P. 245-25I.
ZEDWITZ, F. X. G. (I937). Die Gemse. - Monog. der Wildsaugetiere. III. P. Sch6ps.
Leipzig.
ZUSAMMENFASSUNG
Hornartige Organe haben sich unabhangig voneinander in mehreren Saugerfamilien
entwickelt. So verschieden sie wurden, lassen sie sich doch wenigen Funktionsgruppen
zuordnen.Die Arbeit gibt eine kurze tbersicht uiberBau und Entwicklung hornartiger
Organe und er6rtert kritisch einige Ansichten uiberFunktionenund Stammesentwicklung.
Die Evolution hornartiger Organe lasst sich wie folgt auffassen:
Bei primitivengrossen Saugern wurde das Zuschlagen oder Zustossen mit dem Kopfe
zur wirksamenKampfform, umso mehr, je schwerer die trage Masse des Kopfes. Manche
Formen schlugen diesen Weg ein. Sie kampften breitseits; sie stiessen mit den K6pfen
nach dem Korper des anderen. So wurden Schadelvorspriinge vorteilhaft, und es
entwickeltensich Schutzmechanismen,die die Wirksamkeit dieser Vorspriinge herabsetzten, vor allem eine dicke, schwere Haut oder auf bestimmteOrte beschrankteschildartige
Hautverdickungen.Auf diesem Wege war kein Anlass, besonders grose und spezialisiertere Hornformen zu entwickeln.
Gerade dieser Anreiz aber erwuchs mit dem Auftreten einer neuen Kampfesweise,
bei welcher man die St6sse des Gegners mit den H6rnen auffangt. Das fiihrt zur
Entwicklung schwerer Schadel und H6rner, die den Kopf des anderen einfangen und
festhalten k6nnen. Das Angriffsziel bleibt der K6rper, der Angriff auf ihn wird pariert
durch frontales Gegeneinanderstossender geh6rnten K6pfe.
Rinder- und Schweineartige entwickelten gleichlaufende Kampfesweisen und entsprechendeWaffen. Die Hauer der Eber erfiillen die gleiche Aufgabe und unterlagen
einer ihnlichen Auslese wie die Kopfbewehrungkurzh6rnigerWiederkiuer. So kampfen
Sus und Oreamnos gut miteinander vergleichbar und sind entsprechend funktionel
ahnlich bewaffnet; das gleiche gilt fur Bos und Phacochoerus.
Der urtiimliche Angriff Stirn gegen Stirn fiihrte zu zwei Kampfsweisen, zum
Sichrammenund ,,Ringen".Die ,,Ringer" entwickelten Hornformen, die besonders dazu
geeignet waren, sich in die des Gegners zu verhakeln,so dass sich beide mit voller Kraft
2I4
GEIST, EVOLUTION OF HORN-LIKE
ORGANS
gegeneinander stemmen und miteinander ringen konnten. Die ,Rammer" entwickelten

schwere Schadel und derbe Horner.
Bei Wiederkauernund Schweinen lasst sich eine konvergente Entwicklung besonders
schauwirksamer Gestaltungen des Kopfes feststellen; sie lassen sich als Blickfiinger
der Aufmerkamkeit deuten: ihre Verteilung passt meistens zur Art des Imponierens.
Wiederkauermit gleichmassig verteilten schauwirksamenOrganen imponierenbreitseits.
Wenn die Drohorgane sich mehr auf den Kopf beschranken,verschwindet das Breitseitsimponieren. Das gilt fur die noramerikanischenWildschafe, deren Widder vor
allem mit ihren schweren ausserordentlichlangen Hornern imponieren, die geradezu
als Rangabzeichenwirksam sind. Widder konnen Unterschiede in Horngrosze erkennen.
Je langer und schwerer die Horner, umso dominanter der Widder, umso h6heren
Kampwert traut man ihm zu, und umso mehr Geissen beschlagt er.
So haben sich die H6rner zu vierfacher Leistung entwickelt: als Waffen im Beschadigungskampf, als Verteidigungsschild seines Besitzers, als Klammer, die die
Kampfer fest zusammenschliesst,wenn sie im frontalen Gegeneinanderdrangend ihre
Krafte messen, und als Imponierorganezum EinschiuchternrivalisierenderArtgenossen.
Die Horner, Gehorne und Geweihe mancher Wiederkauerwachsen lebenslang. Offenbar haben sie wie beim amerikanischenWildschaf die gleiche Ranghohen anzeigende
Wirkung. Die Artgenossen lernen, von der Horn- bzw. Geweihgrosse auf den Kampfwert des Rivalen zu schliessen. Bei den grossen Hirscharten gemassigter und kalter
Zonen wird auf diese Art wahrend der Brunst Fett gespart, die den Hirschen hilft,
den strengen Winter zu uiberstehen.Die Verschiedenheitder Horngrosse gestattet den
Wildschafen, in einer ,,offenen" Gesellschaft zu leben, in deren Rangordnungsich frenmde
Hinzukommlingenahezu ohne Kampf einreihen konnen. Bei territorialensozialen Arten,
deren Horner nicht auffallig grossen verschieden sind, dient das Territorium als ein
soziale Verhaltnisse voraussagendesRangabzeichen.
Es wird versucht, die auslesewirksamen Faktoren in der Stammesgeschichte des
Imponierverhaltensaufzuweisen. Es lasst sich voraussagen, dass antagonistische Selektion die Evolution schauwirksamer Organe, Kampf- und Verteidigungsmechanismus,
forderte, und dass ein Selektionsvorteilauf der Geschwindigkeitdieser Parallelentwicklung liegt. Alle Sinnesmodalitateneiner Tierart konnen bei der Herausbildung von
Imponierweisen angesprochen werden, und aussere Erscheinungsmerkmaledes Tieres
konnten sich rascher ahndern, als Anpassungen an das Biotop der Art.
ADDENDUM
One important paper appeared after this paper had been written. FRXDRICH, (i965)
Zs. Tierpsych. 22 (P. 328) reported on the behaviour of several suids including the
warthog. He confirmed and extended MOHR'S(I960) and EWER'S (I958) observationson
the frontal engagement of warthogs. He showed that Sus and Phacochoeruspossess a
broadsidedisplay and are hence similar in this respect to their bovid counterpartsOreamnos and Bos respectively. The suids, while displaying, differ from the bovids only in
that they turn their head towards the opponent rather than away. FRXDRICH states
that DEFFWEILER & MurLER (1924), on the basis of paleontologicalevidence, reached
the conclusion that suid tusks and bovid horns were analogous. Hence, a conclusion
reached independently by EWER (0958) and myself from behaviour data, finds confirmation in entirely different data. FRXDRICH earlier expressed the view that warthog
tusks were not analogous to bovid horns. However, after a discussion of the subject
by mail, he has expressed the same view I propose here.
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The Evolution of Horn-Like Organs

Author(s): Valerius Geist

THE EVOLUTION OF HORN-LIKE ORGANS

studies have been illuminating. It has become evident that the

Eobasileus and the ruminant Protoceras (Fig. I. a, b), or Tetraceros - a

Very little is known of the development and growth-regulating mechanisms

EVOLUTION OF HORN-LIKE ORGANS

VIEWS ON HORN FUNCTION

There is no lack of opinion on horn function and evolution. BRUHIN

I. Horns, similar in shape and size in both sexes function mainly as

EVOLUTION OF HORN-LIKE ORGANS

It is noticeable that horn-like organs appeared and evolved primarily

head swung in an arc or a frontal butt would deliver a substantial amount

EVOLUTION OF HORN-LIKE ORGANS

very widely distributedin mammals.In some, however,it is found only as a

(TEMBROCK, I962), bovids (WALTHER, I958;

observationson Citelluslateralis).The behaviourof many mammalshas not

organs, to make the shield less effective. Equally, there is an impetus to

OF LARGE HORNS AND TUSKS

I have suggested a number of defense mechanisms against the blow of a

EVOLUTION OF HORN-LIKE ORGANS

(1958, I962) on the basis of very extensive observations on

TPwo African elephants (Loxodonta) during an engagement. Note the placement

Observation of Artiodactyla reveal a trend toward cephalization of the

othersthey are cepalsdOrea

(Fig. 5) or of the plain head architecture of Sus scrofa in contrast to the

EVOLUTION OF HORN-LIKE ORGANS

was observed repeatedly by WALTHER (I963, personal communication) in

EVOLUTION OF HORN-LIKE ORGANS

observations suggested that the horns functioned also as display organs,

Ordinateequalspercentfrequencyof sexual approaches.

the next smallersize class respectively.

Fig. 4. During vicious fight (occurring mainly around the estrous

Fig. 7. The low stretch or horn display of sheep, here illustrated

Fig. 8. The horn threat is shown typically by a smaller towards

Fig. 9. Ram bands are assemblies of individuals, three years of ag

larger size classes

smaller size classes

EVOLUTION OF HORN-LIKE ORGANS

Withdrawaltendenciesof rams (Ovis dalli stonei; I962 data)

No. of interactionsin which one ram

degree of dominance, in an association of strange individuals. Ram bands

(a) L Mountswhichledto copulation

Fig. I4. Ordinateequalsmountsperformedby ramson oestrousewes per hourspentin

match allowing test of opponent'sbody strength. Broadsidedisplay prominent.The second

HORNLIKE ORGANS AS RANK INDICATORS AND THEIR

EVOLUTIONOF HORN-LIKE ORGANS

establish such priorities in groups of individuals. An individual without self

EVOLUTION OF HORN-LIKE ORGANS

less effective. If A displays, it is not advantageous for B to withdraw at

bind opponentstogether,thus allowingthem to developfull strengthwrestlingand

that the distributionof displayorgans correlateswith the display.A broadsidebody

Inst. des Parcs Nat. du Congo et du Ruanda-Urundi,Bruxelles.

Theriologicum. Czech. Academy Sci. p. 59-66.

E. H. (I955). The evolution of the vertebrates. COLBERT,

J. Wiley and Sons.

COLLIAS,N. (1956). The analysis of socialization in sheep and goats.

DARLING, F. F. (I937). A herd of red deer. Oxford Univ.

I. (i963). Angeborenes und Erworbenes im Verhalten der Sauger.

(1i96ib). Einige Verhaltensweisen im Hirschrudel. - Rev. Suisse. Zool. ig6i,

GEIST, EVOLUTION OF HORN-LIKE

gegeneinander stemmen und miteinander ringen konnten. Die ,Rammer" entwickelten