The Ecology of the European Roe
Deer (Capreolus capreolus)
Roe deer are one of only two species that can
be considered indigenous to the British Isles, the other being
the Red deer. Remains identified as Roe have been found dating
back to the Interglacial period (400,000 BC) along with other
species now extinct in Britain. There is only one species of Roe
deer, believed to be divided into three subspecies. Considerable
debate exists amongst the experts regarding the exact differences
between the subspecies; this has been compounded by the fact that
the majority of the world’s Roe population lives within the former
Eastern Bloc. As travel restrictions ease, further detailed research
should clarify the issue. The three subspecies are European, Siberian
and Chinese. This paper is primarily concerned with the European
Roe, Capreolus capreolus.
There are significant differences between the
subspecies. The Siberian Roe are approximately twice the size
of European Roe. The latter stand between 60-75 cm at the shoulder
with bucks weighing between 24 and 30 kg, whilst the does are
2-6 kg lighter. Chinese Roe fall somewhere in between these physical
ends of the spectrum. Even amongst the same subspecies there can
be a marked difference in body size, attributable to availability
of food and climatic conditions, the two principal factors which
determine the carrying capacity of a particular habitat. Antler
development varies between the subspecies. Coronets of European
Roe bucks sometimes coalesce. In Siberian and Chinese Roe the
coronets are usually well separated. However, in the Siberian
subspecies the back tines tend to develop a fork and the overall
antler length is greater than that of the Chinese, with the shortest
antlers belonging to European Roe. All subspecies grow the characteristic
six-point head, with minor variations depending on the health
of the deer.
HISTORY AND DISTRIBUTION
European Roe can be considered to have been
continuously resident on the British mainland since the Postglacial
period, crossing from Europe by means of the land bridge. Roe
deer didn't reach Ireland at this time. With the retreat of the
ice pack and the formation of the English Channel they became
marooned on what is now mainland Britain. Since then the native
Roe population has undergone dramatic fluctuations in numbers
Numerous through Roman and Saxon times, Roe
suffered a steady decline through the mediaeval period. Having
been protected by the Normans, Roe were later declared as being
‘beasts of the warren’ (unworthy of noble hunting) in 1338. This
led to their becoming a food source to an expanding peasant population.
By the late sixteenth century Roe were becoming increasingly scarce
throughout England and were extinct in Wales. Scotland experienced
a similar decline in numbers due in part to forest clearance and
the rise of widespread sheep farming. A few small pockets survived
in the southern counties of Scotland, with the remaining population
confined to Highlands.
With the growing interest in forestry and game
shooting during the eighteenth and nineteenth centuries, the fortunes
of Roe deer started to improve. A series of introductions into
parks and wildlife collections in the Lake District, East Anglia
and the South of England re-established the species in the early
nineteenth century. It is worth noting the theory that the Roe
of the Petworth Park Estate were never extinct and the present
population evolved from animals contained within the wall. From
these introductions, escapees and deliberate releases established
wild colonies. Since then there has been a continuing spread across
Britain and, aided by post war afforestation, Scottish Roe have
now closed the distribution gap. Roe can now be found in all counties
in England with a slow but recorded infiltration into Wales. The
attempted introductions of Roe in Ireland, where there has never
been any indigenous population, have met with limited success
and Roe can be considered extremely rare if not non-existent across
the Irish Sea.
Young Roe, born between late May and early
June, are called kids or sometimes fawns. At birth, Roe kids have
a dark brown coat which is heavily spotted with two distinct lines
of white spots running from the nape of the neck either side of
the dorsal to the rump. The sides and flanks are also spotted
in white. The spots soon begin to fade and are usually gone in
about 8-10 weeks. The kids then develop their first winter coat.
By September/October Roe are in full winter pelage. Their coat
colour ranges from dark brown to charcoal grey. The rump hairs
or caudal patch, erectile when the animal is alarmed, is white
and far more prominent in winter pelage. The caudal patch is important
in identifying the sexes during the winter when the bucks are
re-growing their antlers. The does have a distinctive downward
pointing tuft of hair at the base of the caudal patch referred
to as an anal tush, which gives the appearance of an ace of spades.
Bucks do not have this distinguishing feature and their caudal
patch resembles a kidney shape. The winter coat is shed from April
to May, younger animals changing first. Whilst the winter coat
is being shed, Roe take on a moth-eaten and unkempt appearance
as their coat falls out in chunks, usually starting around the
neck, revealing the summer coat underneath.
The summer coat of both male and female is chestnut red although
it can vary to a sandy yellow. Occasionally an animal with a darker,
black or piebald pelage is seen although this is uncommon. It
is often the unfortunate individual with a different or unusual
pelage that ends up being taken as a ‘unique’ sporting trophy.
Roe have a very short tail, which is not visible
without close physical examination and is an important distinguishing
feature of the species. Both bucks and does are marked on the
face with a white spot on the upper lip either side of the nose
and the chin is also white. This coloration is more prominent
in younger animals.
As ageing takes place, an increasing amount of grey develops
around the muzzle and the hair on the forehead becomes curly,
giving bucks an ‘old man’ appearance. The study of variegation
(different colouring) of a Roe’s head and face can be used as
an aid to ageing under field conditions.
Roe deer are unusual amongst British deer in
that they cast and re-grow their antlers in the winter when good
food is scarce; other species develop their antlers over spring
and summer. Buck kids develop a small knob (or button) on top
of their pedicle during the first nine months of life. This knob
is shed in early spring and the first antler begins to grow. Occasionally
if habitat conditions have been favourable, a strong young buck
may develop a classic six-point head in its first year. However,
a four-point head or even just two spikes would be the norm.
During its development, the growing antler
is covered in a grey furry membrane known as velvet. This carries
the blood vessels and nerves for the developing antler; the antler
is bone growing on the outside of the body from the pedicles unlike
horns, which are hollow and not cast each year. Should the velvet
become damaged during the antlers’ growth, deformities can occur.
The antlers are usually fully developed between March and April
(earlier in older animals), when the increasing length of daylight
causes a rise in testosterone levels. This causes the blood supply
to the antler to be cut off. The velvet dries out, shrinks and
is removed by the buck fraying its new antlers against saplings.
During the period that the velvet is being removed, the buck is
said to be ‘in tatters’ with dried velvet hanging in strips from
the now hard antler. Whilst removing the velvet, damage is caused
to young trees as the buck frays. It is whilst the buck is fraying
saplings that the antlers become stained and coloured with tannin
and sap from the bark, which changes the colour from white bone
to a deep burnished brown. In areas with few trees like Salisbury
Plain, antler colour is much paler due to lack of bark staining.
It is the older bucks who clean their antlers first in readiness
From April onwards mature bucks begin to mark out their territories.
Trees are now frayed for a different reason. This is boundary
scent marking; scent from the preorbital gland is deposited onto
the wet sapwood under the bark. This second fraying causes significantly
more damage than the cleaning of velvet. It is considered good
management policy to leave a territorial buck to dominate a new
plantation and act as a policeman against questing bucks seeking
territories. Removal of a stand buck often results in an influx
of younger males resulting in much frustrated fraying and considerable
damage. Frayed herbage normally has a V-shaped scrape at its base
which is made by the buck with its front hooves. This will also
be scent-marked by secretions from the interdigital glands situated
between the cleaves of the deer’s hooves.
Animals still in velvet in June may well have
been injured or might be unwell for some reason. There is a condition
in Roe which can lead to the development of what is known as a
Perruque Head. This is normally caused by damage to a buck’s testicles
which prevents the rise in testosterone which would normally signal
the cessation of the blood supply to the antlers’ velvet. Consequently
with a perruque the velvet continues to grow, forming a soft spongy
mass which does not harden. This eventually becomes infected or
fly blown and may result in the animal’s death.
The typical European Roe
buck grows two, three point antlers consisting of a brow point,
a top point and a back point. The base of the antler is called
the coronet or burr and grows from the pedicle. The coronet increases
with size as the animal ages, sometimes growing together (coalescing)
in older animals. The beam of the antler is covered in tiny bony
nodules called pearling. Antler development in Roe is dependent
on food supply and age. A mature animal in poor condition will
only produce a small head. If conditions are better the following
year then a fine head may be grown. The average length for European
Roe antlers is between 20-30cm. The bucks shed their antlers between
October and December, the older bucks shedding first. New growth
starts almost immediately. Older bucks can be identified from
their antlers as the coronets begin to droop (like melting candle
wax) and they angle forward losing their crisp right angle to
the pedicle. The tines lose their well developed shape, becoming
less sharp, and sometimes there is slight palmation at the joint
of tine and beam. A buck in this condition is termed as ‘going
back’. Occasionally very old does will develop pedicles and sometimes
small antlers; this is an hormonal imbalance due to old age.
HABITAT AND FEEDING
Roe are herbivores and a highly adaptable species
found in a wide variety of habitats, ranging from open moor to
thick cover in conifer or deciduous woodland. Features that must
be present whatever the habitat are shelter and a variety of food
plants. The ideal habitat could be considered as coppice and pockets
of deciduous woodland on land that is not intensely farmed, this
linked by thick hedgerows and dotted with small copses. Roe are
browsers, however they also graze, feeding on shoots, herbs, grasses,
fruits, nuts, fungi, pine needles and twiggy browse during hard
times. Roe are opportunistic feeders with a liking for the exotic
as many gardeners have discovered. They can decimate rose beds
and low hanging baskets as well as vegetable gardens given the
chance. The feeding cycle of Roe is the same as with other British
deer. Being ruminants they feed in bouts followed by periods lying
up to chew regurgitated cud. However the digestive system of the
Roe is simpler than that of other species and this causes food
to pass through the alimentary tract relatively quickly. In turn
this necessitates more frequent feeding bouts depending upon the
quality and quantity of food available. Should the animals feel
secure then it is not unusual for them to lie where they have
Roe feed heavily during the spring and summer
months, building up reserves to see them over the winter - the
bucks will have new antlers to grow and most does will have a
pregnancy to carry to full term. During late autumn and winter,
activity decreases due to a reduced metabolic rate (a form of
semi-hibernation). If not disturbed they can refrain from feeding
for longer periods than normal. If disturbed then food will be
required to replenish expended energy and poor food quality at
this time of year necessitates greater intake in order to extract
Roe can be seen feeding at all times of the
day depending on season and weather. The main times of activity
are at dawn and dusk (crepuscular). If regularly disturbed they
can become nocturnal, staying in close cover during daylight.
Roe feeding in the open will head directly for cover if disturbed.
Heavy rain and strong winds tend to suppress feeding activity
and they will stay in cover. So a good time to find Roe feeding
is after inclement weather as they quickly emerge to feed and
dry off. Roe like to sunbathe and a sunlit clearing in cover will
often be a favourite place to lie up.
The type of soil in an area plays a major roll
in the expected lifespan of Roe. A deer lives as long as it is
able to chew its food enough to extract essential nutrients. In
areas of harsh gritty soil teeth wear away more quickly than in
areas of soft loamy soil, the sand and grit acting as an abrasive
on the teeth. Tooth wear is location specific and with local knowledge,
wear is a good guide to a deer’s age. In young animals very little
wear will be seen, but in very old animals the teeth may be worn
down to gum level and like this the animal will eventually die
from malnutrition. An accurate way of assessing the age of an
animal in its first year is by examination of the third pre-molar
tooth. At birth this milk tooth is divided into three crowns.
This tooth is replaced by the permanent 2 crown pre-molar by the
time the animal is 13 months of age and it will generally have
less staining than the other teeth.
Roe in Britain are not normally gregarious
animals; hence they are not farmed. They are normally seen in
loose family groups or as single animals. However in the Salisbury
Plain area they mimic continental Field Roe in that in winter
they can be seen in groups of up to 14 animals. The majority of
a buck’s life is solitary. The doe will be accompanied by kids
for most of the year and when she appears alone it is likely that
there will a buck not too far away. Spring is a time of much activity.
Individual bucks set up or re-establish territories where possible,
however not all manage to achieve this. Bucks without status are
pushed to the periphery of good areas where food and cover are
less favourable. It is this action which leads to emigration and
immigration between areas and is very significant in relation
to Roe distribution. The does also drive the previous year’s kids
from their home range in readiness to give birth to this year’s
offspring. Many animals, having been ousted from their familiar
surroundings, wander far afield seeking a favourable place to
live. This results in an increase in road traffic accidents at
this time of year. Roe will often congregate in an area not dominated
by territorial bucks or does, known as a ‘coffee bar’ or more
accurately, a ’non-disputed area’ where they can cause considerable
Occasionally a weak young buck who is not considered
a threat by a stand buck, will be tolerated within a defined territory.
This affords the young buck the benefit of good cover and food,
enabling it to develop in physical stature and establish a knowledge
of the area. Known as a Satellite buck, the young male risks being
frequently chased and even injured by the resident buck when he
attempts to court a female. It is very likely that this animal
will eventually inherit the territory.
Peripheral bucks reside in the core areas of
the buffer zones between territories and make short raids into
established territories in search of unattended females. If a
buck has not established a territory by its third birthday, it
is unlikely to ever achieve one.
The does remain in their home range or territory
for most of the year as this favours their offspring. A doe does
not make physical boundary markings like the buck’s fraying, but
will mark her territory with scent from the interdigital glands.
Other does and eventually her own kids will be driven off, sometimes
with physical force using the front hooves in a rearing and stabbing
motion. A doe’s range may overlap that of other does and these
will probably all overlap a buck’s territory.
The majority of Roe kids
are born in late May and early June, although kids have
been seen as early as March and as late as August in the South
of England. Roe are unique amongst British deer in that the process
of fertilisation after mating is subject to a biological mechanism
called embryonic diapause or delayed
implantation, as are other British mammals such as badgers
The rut takes place between late July and early
August, but after fertilisation the embryo continues to float
free in the uterus developing very slowly. It is not until late
December or early January that the embryo becomes implanted in
the uterine wall and rapid fetal development begins. Recent scientific
research by several universities across Europe have shown female
roe deer to have a single annual sexual cycle with a well-defined
breeding season (they are described as monoestrous). During her one reproductive cycle,
a doe sheds eggs from her ovaries, which are fertilised when they
are in the fallopian tubes (the narrow tube between the ovaries
and the uterus). Over the next 24 hours the embryo begins cell
division as it passes into the uterus. When the developing embryo
reaches 20 – 30 cell stage (known as a blastocyst) it enters a
5-month period where it remains unattached to the uterus. This
period is known as delayed implantation or embryonic diapause.
Left behind in the ovaries is the corpus luteum,
one for each egg shed and are only a few millimetres in diameter.
The corpora lutea release progesterone, which provides the embryo
with nutrients to maintain the pregnancy. In species, other than
the roe deer, the corpora lutea are normally an indication of
pregnancy because they regress if fertilisation has not occurred.
If the eggs fail to be fertilised in the roe doe, the corpora
lutea in the non pregnant doe remain active for a period of five
months, which coincides with the period of delayed implantation
in the pregnant doe. This period in the non-pregnant doe is known
as pseudo-pregnancy. Roe are one of only two species of mammal
to have active corpora lutea during the period of delayed implantation,
the other species being the armadillo. This means that the number
of corpora lutea present in the ovaries during the period of delayed
implantation do not give an accurate guide as to the number of
embryos she might have or whether she is in fact even pregnant.
During the period of delayed implantation the doe cannot know
whether she is pregnant or not, because the embryos, during the
delay period, do not send any 'signals' to the mother that they
are there, hence she keeps the corpora lutea active whether she
has embryos or not. She is in fact 'assuming' she is pregnant.
The roe mother only recognises pregnancy when the fetus implants
in the wall of the uterus in January. Therefore, if no embryos
implant, the doe knows she is not pregnant and the corpora lutea
start to regress quite rapidly from January onwards, whereas,
in the pregnant doe, they remain active until the birth of the
kids in late May. No other species of deer is monoestrus
or employs delayed implantation, including the Siberian roe, as
a reproductive strategy. Other deer species are described as polyoestrous
and have many sexual cycles, usually occurring in the autumn.
The breeding potential of a doe is dependant upon her body condition
at the beginning of the rut. An animal that has wintered in good
conditions will come into oestrus sooner than a doe that has had
to cope with a harsher environment. Yearling does (maidens) come
into oestrus a little earlier than mature does and are often covered
by bucks before the main rut begins. The doe can be considered
the more sexually ‘dominant’ of the sexes. If there is not a buck in the area when a doe comes
into oestrus, she will go and find one and will then lead it to
her preferred area with much leading on and chasing. Once satisfied
that the buck is strong and worthy of passing on its genes, copulation
will be permitted. The doe may be covered many times over a short
period. Should the doe not be fertilised by the first attempts
at mating, then she will come into season again a few days later
and be covered by the same or another buck. Contests over available
does by bucks can be ferocious and prolonged affairs. If the initial
challenging bark fails, head on fights occur until one concedes
that the other is the stronger by retiring. Injuries are not uncommon,
indeed they can sometimes prove fatal. Normally facial and neck
wounds can be evident, however if the vanquished buck is slow
to leave then puncture wounds to the hindquarters may be inflicted.
It is in the rut that bucks are at their most
pugnacious and inquisitive. They will approach a detected presence
and even respond to an approximation of the call of a doe in oestrus,
or the ‘pheep’ of a kid which may indicate that a doe is near
by. A buck will approach expressing a challenge by stamping its
foot, raising and lowering its head, licking its nose to improve
its scenting ability and giving a challenging bark. Only when
the buck has caught a human scent or recognised a threat will
it bound off. However, another imitation call at this juncture
is often enough to stop it in its tracks for a second look.
In good conditions Roe have
a high fecundity rate. In the South of England twins are the norm
and in very good conditions triplets are not uncommon. Many triplet
fetuses are found during a post-mortem examination, though not
all would necessarily have been born alive. But if all three fetuses
are the same size, the indications are that all three will be
live at birth. Should one or more be smaller than the others,
then it is unlikely it would have been born alive. In a particularly
cold and/or wet early summer and autumn, the survival of all three
triplets is greatly reduced and it is normally buck kids that
fare worst. This could be due to nature perpetuating the species
by saving the does for the following breeding season.
VOICE AND CALLS
The main call of the Roe is a bark. The buck’s
bark is usually a challenge call; the bark resembles that
of a medium size dog, the timbre of the call becoming deeper as
the animal ages. Bucks make a variety of other noises during the
rut when pursuing a doe. However, the doe is the most vocal of
the sexes and can be heard barking at most times of the year.
The doe mostly barks as a warning to her kids and may continue
to bark in various patterns for some time. Other calls consist
of ‘pheeps’ and squeaks when communicating with her kids. The
doe has a definite call when in oestrus and this is used to advertise
her condition to any nearby bucks.
Whilst Roe have a remarkable sense of smell,
their eyesight is limited as they have monochromic vision, seeing
in shades of black and white. They can detect light reflection
from an unnatural material and it is this rather than colour that
alerts Roe to danger. However Roe are able to detect the slightest
physical movement and can spot a moving human form at considerable
Roe are probably the most fascinating and interesting
species of British wild deer. They are also the most rewarding
species to watch as, apart from the challenge of getting close
enough to observe them, their characteristics and behaviour are
thoroughly individualistic. Their many unique features, along
with a natural elegance and grace, place them high on the list
of our nation’s natural assets. With the planned expansion of
the National Forest over the next twenty years, Roe will take
full advantage in exploiting new habitats. The present Roe deer
population is probably at its highest since the Middle Ages.
Richard Prior, The Roe Deer- Conservation of
a Native Species. Swan-Hill Press 1995.
Richard Prior, Roe Stalking. Game Conservancy
Richard Prior, Deer Watch. Swan Hill Press
Ian Alcock, Chasing the Red and Following the
Roe. Sauchenyard Press 1998.
The British Deer Society, Basic Deer Management.
Game Conservancy Publishing, Deer Management
in Small Woodlands.
Kenneth Whitehead, Practical Deer Stalking.
AJ de Nahlik, Management of Deer and Their
Habitat. Wilson Hunt 1992.
Herbert Krebbs (translation by Lt Col Roy C
Harms HQ USAREUR), Young or Old. F.C. Mayer Munchen-Solln.
Liberg, O., Johansson, R., Andersen, R. and
Linnell, J.D.C. (1998) The Function of Male Territoriality in
Roe Deer. In: The European Roe deer: The Biology of Success,
Scandinavian University Press UK, Oxford.
Additional information supplied by Roger Lambert.
Lambert, R.T., Ashworth, C.J.,
Beattie, L., Gebbie, F.E., Hutchinson, J.S.M., Kyle, D.J. &
Racey, P.A. 1999. Temporal changes in reproductive hormones and
conceptus-endometrial interactions during delayed implantation
in the European roe deer (Capreolus capreolus). In: Advances
in Deer Biology (Ed: Z. Zomborszky), Pannon University, Hungary.
The Deer-UK Team are grateful to the University
of Aberdeen for allowing the use of the Roe buck photograph in
the Antler Development section.