Animal Reproduction Science 124 (2011) 170–175

Contents lists available at ScienceDirect

Animal Reproduction Science

journal homepage: www.elsevier.com/locate/anireprosci

Reproductive cycles of deer夽

G.W. Asher
AgResearch Ltd., Invermay Agricultural Centre, Private Bag 50034, Mosgiel, New Zealand

a r t i c l e i n f o a b s t r a c t

Article history: The cervids are a complex assemblage of taxa showing extreme diversity in morphol-
Available online 29 September 2010 ogy, physiology, ecology and geographical distribution. Reproductive strategies adopted
by various species are also diverse, and include a range from highly seasonal to com-
Keywords: pletely aseasonal birth patterns. The recent growth in knowledge on cervid reproduction is
Deer strongly biased towards the larger-bodied, gregarious mixed grazer-browser species that
Reproductive cycle
have adapted well to human management and commercialisation. These species tend to
Pregnancy
represent ‘K-selected’ climax species characterised by very productive annual breeding
Seasonality
success, singleton births and long breeding life (10+ years). Conversely, we know relatively
little about the reproductive patterns of the ‘r-selected’ smaller-bodied, solitary (and often
highly territorial), forest-dwelling browser species, often characterised by great fecundity
(twinning) and shorter breeding life (<10 years). This group includes many of the endan-
gered cervid taxa. This review extends earlier reviews to include more recent work on cervid
reproductive cycles, particularly in relation to environmental factors influencing gestation
length.
© 2010 Elsevier B.V. All rights reserved.

1. Distribution and diversity of cervids
The ‘cervids’, represented by over 40 species and
200 subspecies (Whitehead, 1993), are characterised by
remarkable diversity in morphology, physiology, ecology
and geographical distribution. In particular, reproductive
strategies amongst cervids vary widely within and between
regions. As such, no one species can be considered to rep-
resent a ‘typical’ deer in terms of reproductive function.
For example, some cervids exhibit highly seasonal pat-
terns of birth in cool temperate climes, while others are
aseasonal in equatorial regions. Also, while many species
are strictly monovular and bear single offspring annu-
ally, others are normally polyovular and bear multiple
offspring. Even embryonic development and placentation
vary amongst species (Asher et al., 1999a,b). Harrington
(1985) expressed this variation in terms of ‘life-history
夽 This paper is part of the special issue entitled: Reproductive Cycles of
Animals, Guest Edited by Michael G. Diskin and Alexander Evans.
E-mail address: geoff.asher@agresearch.co.nz.
strategies’ in which cervids can be classified into the two
extremes; one maximises the reproductive capacity of indi-
viduals and the potential growth rate of the population
(r), and the other maximises the competitive ability of
individuals and the stability of the population (K). Unsta-
ble or early-successional habitats favour the r-strategy,
while stable or late-successional (climax) habitats favour
the K-strategy (Harper et al., 1961). The r-strategists are
identified by their greater reproductive potential, exhibit-
ing multiple births, early sexual maturation (<1 year) and
short breeding life (<10 years). They are predominantly ter-
ritorial browser- or concentrate-feeders adapted to rapid
colonisation of early stage successional habitats; including
species such as roe deer (Capreolus capreolus), white-
tailed deer (Odocoileus virginianus), moose (Alces alces)
and numerous species of tropical, forest-dwelling deer. K-
strategists have lesser reproductive potential that includes
lower fecundity (i.e. singleton births) and later sexual mat-
uration (>1 year) and are generally mixed grazer/browsers
with marked gregariousness and social/behavioural plas-
ticity. They tend to occupy climax habitats and include
nearly all taxa within Cervus (red deer, wapiti, sika deer,

0378-4320/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2010.08.026
 G.W. Asher / Animal Reproduction Science 124 (2011) 170–175
sambar deer, etc.) and Rangifer (reindeer, caribou). Because
of their ability to adapt to human management regimens,
it is the K-strategists that are predominantly represented
in the research literature. It is therefore important to note
that while there was an increased understanding of repro-
ductive function in cervids over the last few decades, this
relates mainly to those species of importance to agri-
culture (i.e. deer farming) and a few endangered species
undergoing ex situ conservation efforts. In general, the
research has been largely directed towards successful
application of assisted reproductive technologies to manip-
ulate the genetic composition of populations (e.g. selection
for improved agricultural productivity and avoidance of
inbreeding depression). The studies to date are clearly
biased towards a few key species, particularly the larger-
bodied, monovulatory species with greater environmental
and behavioural plasticity (K-strategists) that makes them
suitable candidates for captive propagation. The smaller-
bodied, forest-dwelling and solitary species (r-strategists)
are clearly under-represented in such research. This later
group includes numerous species whose populations are
under threat or are endangered (see Asher et al., 1999a).
2. Seasonality compared to aseasonality
The necessity for most cervid species to give birth at an
appropriate time of year for optimal survival and growth
of offspring has exerted considerable influence on their
reproductive physiology (Lincoln and Short, 1980). Species
of northern temperate or arctic origins typically conceive
in autumn and calve in summer. In contrast, species of
tropical origin often exhibit limited seasonality or are
completely aseasonal (Lincoln, 1985). The endogenous
mechanisms governing seasonal reproductive patterns in
temperate species are robust, being manifest rigorously
when animals are transferred between localities despite
subtle variations in seasonal feed supply. The transfer-
ence of temperate species across the equator results in
an exact 6-month phase change (Marshall, 1937), even
though the relationship between season and feed pro-
duction differs considerably between continental northern
hemisphere and insular southern hemisphere environ-
ments (Asher et al., 1993a). Tropical species transferred
from equatorial zones exhibit either pronounced ‘reverse
seasonality’ (spring conceptions and autumn/winter calv-
ing; e.g. rusa deer, Cervus timorensis, in southern Australia),
wide birth seasons (e.g. chital deer, Axis axis, in Australia)
or complete aseasonality (e.g. Reeve’s muntjac, Muntiacus
reevsii, in UK) (Chapman et al., 1984; Loudon and Curlewis,
1988; Monfort et al., 1991; Mylrea, 1992). It is notewor-
thy that there is a general paucity of information on the
birth season of tropical species in their native tropical
environment, although there are well-documented exam-
ples of species that exhibit strictly seasonal patterns of
birth.
2.1. Photoperiodic control of reproduction
For temperate and arctic species it is generally accepted
that entrainment of seasonal reproductive cycles is effected
by endogenous recognition of photoperiod changes, with
171
the majority of species initiating mating activity during
decreasing daylength of late summer and autumn (Lincoln
and Short, 1980). Variations in the actual mating season
between species of up to 8 weeks usually offsets differences
in the duration of gestation, such that parturition generally
occurs in mid-summer (Lincoln, 1985). A notable exception
amongst temperate species is the Pere David’s deer (Ela-
phurus davidianus), which initiates mating activity in early
summer and calves in late spring despite an unusually long
gestation of ∼280 days (Wemmer et al., 1989).
There is increasing evidence that some tropical species
also perceive, and respond to, changes in photoperiod,
although in a different manner than temperate species.
Rusa deer living in the tropics exhibit asynchronous antler
cycles and birth timings. However, once translocated to
temperate, high-latitude zones they appear to establish an
evolutionary-based ‘long-day’ reproductive rhythm (Van
Mourik and Stelmasiak, 1990). Brow-antlered Eld’s deer
(Cervus eldi thamin) exhibit strong reproductive seasonal-
ity in their sub-tropical native range. This pattern persists
after translocation to high latitudes (48◦ N), even though
this strategy results in the birth of offspring during harsh
winter months (Monfort et al., 1991). It is hypothesised that
Eld’s deer respond to the relatively subtle low-amplitude
photoperiodic oscillations experienced in their native sub-
tropical latitudes. Higher latitudes simply serve to reinforce
or strengthen seasonal rhythms because the direction of
photoperiodic change is similar, just of greater ampli-
tude. Chital deer appear to exhibit only limited seasonality
within their native tropics and following translocation to
temperate zones. However, exogenous melatonin treat-
ment has been shown to hasten and synchronise antler
casting amongst stags (Mylrea, 1992), indicating possible
photoperiodic responsiveness in a species that, in other
respects, appears unresponsive. In contrast, tropical sam-
bar deer (Cervus unicolor) acclimatised to temperate zones
in New Zealand exhibit a loose pattern of ‘reverse season-
ality’ in calving but fail to show any discernable seasonal
patterns in prolactin secretion (Asher et al., 1997a).
Because tropical cervid species are not exposed to
strong circannual photoperiodic rhythms, reproductive
patterns may have evolved in response to a variety of fac-
tors including seasonal fluctuations in the availability of
specific food items, local resource competition amongst
sympatric species (species that occupy about the same area
of land but do not interbreed), or predation pressure; and
many of these factors may be directly related to rainfall pat-
terns. For example, it has been asserted that conceptions
are timed in white-tailed deer living in Columbia (5◦ N) and
the Everglades National Park (25◦ N) so that births occur
during the dry season (Blouch, 1987; Smith et al., 1996). The
fawning season of hog deer (Axis porcinus) in Nepal (27◦ N)
coincides with new vegetative growth that occurs after sea-
sonal grass fires caused by lightning strikes or agricultural
burning, believed to have been occurring for thousands of
years (Mishra and Wemmer, 1987). However, how these
species synchronise their season of conceptions to achieve
birth synchrony remains to be resolved. Even species like
sambar deer and chital deer sharing the same habitat in
Nepal fawn 6 months out-of-phase to one another (Mishra
and Wemmer, 1987).
172 G.W. Asher / Animal Reproduction Science 124 (2011) 170–175
It is this lack of reproductive uniformity amongst sym-
patric species in the tropics and subtropics from which
can infer differing adaptations to the environment based
on a variety of regulatory mechanisms. In Nepal’s Royal
Chitawan National Park four cervid species co-exist, includ-
ing two browsers (sambar deer and muntjac), one grazer
(hog deer) and one mixed feeder (chital deer). Each of these
species differs in their annual reproductive cycle, habitat
selection and feeding ecology (Mishra and Wemmer, 1987).
Although photo-responsiveness in tropical cervid species
may persist as an evolutionary relic, the tight linkage
between photoperiod and seasonal reproduction observed
in temperate species may not completely explain the exis-
tence of reproductive seasonality in many tropical species.
2.2. Subspecies differences (e.g. red deer)
Cervid species occupying wide latitudinal and longitu-
dinal ranges have generally developed discrete populations
that exhibit pronounced phenotypic and genotypic differ-
ences. For example the red deer/wapiti species (Cervus
elaphus) includes at least 23 recognised subspecies in a
west/east cline from Western Europe to North America
(Whitehead, 1993). Climatic variation between subspecies
zones may be associated with subtle variation in reproduc-
tive seasonality of the various subspecies. Recent studies
have indicated that Western European (Scottish) red deer
(C.e. scoticus) and Eastern European red deer (C.e. hippela-
phus) farmed in New Zealand exhibit a 2–3 week difference
in rutting activity and subsequent mean calving dates
(Scott et al., 2006). These differences occur at the same lat-
itudes in both Europe and New Zealand (and indeed within
sympatric populations on New Zealand farms), indicating
subtle genetic differences in subspecies responses to the
prevailing photoperiodic regimen.
3. Female
3.1. Seasonality of the oestrous cycle
With few exceptions, female cervids are polyoestrous,
and non-pregnant animals are capable of exhibiting contin-
uously repeated oestrous cycles (some tropical species) or,
more commonly, alternating periods of oestrous cyclicity
and anoestrus. Oestrous cycles have been characterised for
a number of species from studies on oestrous behaviour
and/or luteal secretion of progesterone. In non-pregnant
females of temperate species, such as red deer and fal-
low deer, the onset and termination of oestrous cyclicity
occur in autumn and spring respectively, with 5–8 cycles
expressed. Anoestrus is characterised by low peripheral
plasma concentrations of progesterone indicative of com-
plete ovulatory arrest, and may persist for 4–6 months
from spring to early autumn. In marked contrast, some
tropical species, such as Brow-antlered Eld’s deer and rusa
deer, translocated to higher, temperate latitudes exhibit
‘reverse seasonality’, initiating oestrous cycles in spring
and entering anoestrum in the following autumn. This 6-
month phase shift in tropical species within temperate
climes possibly indicates retention of photoperiodism from
a temperate ancestral form. Some tropical species, such as
chital deer appear to have become completely aseasonal,
with females exhibiting free-running oestrous cycles in the
absence of pregnancy (Mylrea, 1992). There are indications
for some tropical species within temperate latitudes, such
as sambar deer in New Zealand, that responsiveness to pho-
toperiod has been lost, as indicated by an inability to detect
measurable levels of plasma prolactin in sambar hinds dur-
ing the year (Asher et al., 1997a).
3.1.1. Oestrus and ovulation
The transition into the female breeding season is char-
acterised by ‘silent ovulations’ (ovulations not preceded by
overt oestrus) and short-lived (8–10 days) corpora lutea in
most cervid species studied. In fallow deer (Dama dama),
multiple successive silent ovulations leading up to the start
of the breeding season have been observed (Asher, 1985).
The transient nature of the preliminary corpora lutea may
actually serve to promote within-herd synchrony of first
overt oestrus of the season.
Subsequent oestrous cycles are generally of normal
duration, as determined genetically for each species,
although occasional ‘long cycles’ have been observed in
some Pere David’s deer and Eld’s deer (Curlewis et al.,
1988; Monfort et al., 1991). Average duration of the nor-
mal oestrous cycle ranges from 17 days in sambar deer and
axis deer, 18–20 days in red deer and Eld’s deer, 21–23
days in fallow deer, and 24–27 days in moose and black-
tailed deer (Odocoileus hemionus). The mean duration of
the oestrous cycle tends to increase progressively during
the breeding season in most of these species. (Guinness et
al., 1971; Asher, 1985; Curlewis et al., 1988; Monfort et
al., 1991; Chapple et al., 1993; Schwartz and Hundertmark,
1993; Wong and Parker, 1993).
Overt oestrous behaviour has been observed in a num-
ber of cervid species but the duration of mating receptivity
at a single oestrus has been difficult to determine. In some
species at least, including red deer and fallow deer, overt
oestrus is generally terminated following successful copu-
lation and may only persist for a matter of minutes. In the
absence of copulation, oestrous behaviour in these species
has been observed to occur for up to 24 h (Asher et al.,
1993b). Early studies on both of these species have shown
that the onset of oestrus coincides with the onset of the pre-
ovulatory LH surge, which itself peaks about 24 h before
ovulation (Berg et al., 1994, see Asher et al., 1999a).
3.1.2. Luteal development
Luteal events during the cervid oestrous cycle are
similar between species. Luteinisation of post-ovulatory
follicles is associated with increased secretion of proges-
terone, with maximal concentrations in peripheral blood
occurring between days 10 and 16 of the oestrous cycle
(Day 0 = oestrus). Although absolute plasma concentrations
vary between species, the relative changes from Day 0 are
quite similar between species (Asher et al., 1999a). Luteal
regression, which for fallow deer involves the classic inter-
action between oxytocin and prostaglandin F2a (Asher et
al., 1988a), occurs rapidly about 1–3 days before the return
to basal progesterone concentrations and return oestrus.
Ovarian follicular dynamics during the oestrous cycle
have been described from ultrasonographic studies on
 G.W. Asher / Animal Reproduction Science 124 (2011) 170–175
red deer, fallow deer and, more recently, North Ameri-
can Wapiti (C.e. manitobensis) (Asher et al., 1997b, 1999b;
McCorkell et al., 2006). These studies, all of which involve
monovulatory species, clearly demonstrated discrete, non-
random patterns of antral follicular growth and regression
during the oestrous cycle, with individual cycles charac-
terised by 2–4 dominant follicle waves, similar to that
observed in cattle.
3.1.3. Exogenous manipulation of ovulation
In many respects, the cervid oestrous cycle is similar to
that of domestic ruminant species. Thus, methods of artifi-
cial oestrous synchronisation have tended to follow similar
procedures used for sheep and cattle. These include, prin-
cipally, the use of progestagens to simulate the luteal cycle,
and prostaglandins to control luteal longevity (see review
by Asher et al., 1993b). As a generalisation, these tech-
niques have been efficacious, leading to the development
of a number of successful technologies such as artificial
insemination and embryo transfer. However, apart from
their application to the genetic management of farmed red
deer and fallow deer in New Zealand, such technologies
are not currently adopted widely for cervids (Morrow et
al., 2009).
3.1.4. Pregnancy/gestation
Relatively little research has been dedicated to preg-
nancy establishment and maintenance in cervids. In the
seasonally breeding red deer hind, conception rates fol-
lowing natural mating during the autumn rut appear to
be high (>85% per mated oestrus), and early embryonic
mortality (i.e. within 20 days of fertilisation) appears to be
low (i.e. <1%; Berg et al., 1994). Given that hinds are poly-
oestrus, under circumstances where nutrition and disease
are not limiting, adult female red deer generally exhibit
greater annual fertility (i.e. >96% pregnancy rates), albeit
with lesser (i.e. <1% of pregnancies) twinning (Asher and
Pearse, 2002). This overall pattern of greater annual fertility
appears to be a common feature of many cervids, although
actual fecundity varies between species. As expected for
the greater fecund species, such as white-tailed deer and
roe deer, the actual fecundity (i.e. incidence of multiple-
foetus gestations) in a given year and locality is strongly
influenced by habitat and nutritional quality variables.
Oocyte maturation, fertilisation and early embryonic
development have recently been studied in vivo and in vitro
for red deer (Berg et al., 2008). These studies have shown
that these events in red deer are similar to those occur-
ring in other domestic ruminants, with the exception that
the red deer embryo enters the uterus 2–3 days later at
the blastocyst stage, 5–6.5 days after ovulation. Embryonic
implantation in red deer occurs shortly before day 27 from
ovulation (McMahon et al., 1997).
Luteal support of pregnancy in red deer persist through
the term of gestation, with peripheral blood progesterone
concentrations remaining elevated to within 1–2 days of
parturition. The contributory role of placental progesterone
production in pregnancy maintenance is unknown. Stud-
ies on red deer and white-tailed deer clearly show that
the presence of a functional corpus luteum is a requisite
for a viable pregnancy (Asher et al., 1996; Plotka et al.,
173
1982). However, ovariectomy of reindeer in late preg-
nancy was not always associated with abortion, indicating
a possible contributing role of the foeto-placental unit in
progesterone secretion and pregnancy maintenance in this
species (Sjaastad et al., 1990).
Importantly, these studies may not be representative of
cervids in general. Placentation systems vary across cervid
species (McMahon et al., 1997), and in the case of one
species, the roe deer, a form of embryonic diapause/delayed
implantation occurs to lengthen the interval between mat-
ing and births in order to ensure optimal birth seasonality
(Short and Hay, 1966). This is the only ruminant species
known to exhibit delayed implantation.
3.1.5. Genetic compared to environmental control of
gestation length
Cervid species have gestation lengths ranging from
200 to 280 days, loosely correlated with species mature
live weight. There is a general underlying acceptance that
gestation length for any given species is genetically pro-
grammed and relatively robust in the face of various
environmental conditions. Recognition is given to the influ-
ence of foetal genotype as being the single most important
determinant of gestation length: as well as determining
interspecific differences in gestation length, it also accounts
for subspecies and breed differences (Kenneth and Ritchie,
1953; Racey, 1981). Indeed, this has been demonstrated for
the red deer species (Cervus elaphus) in which there is con-
siderable subspecies variation in observed mating-birth
intervals ranging from 233 days for Scottish red deer (C.e.
scoticus) to 247 days for North American Wapiti (C.e. nel-
soni, roosevelti, manitobensis) (Guinness et al., 1971; Haigh,
2001). Red deer hinds gestating F1 crossbred red × Wapiti
foetuses show a mating-calving interval of ∼239 days
(Asher et al., 2005b). Similarly, although Pere David’s deer,
with a reported gestation length of ∼280 days, are classified
within a different genus, hybridisation with red deer hinds
can result in fertile offspring. Hinds gestating F1 hybrid
red × Pere David foetuses exhibit a wide range in mating-
calving intervals from 262 to 274 days (Asher et al., 1988b).
However, it has become apparent that gestation length
within red deer subspecies is much more variable than pre-
viously thought (Asher, 2007). Guinness et al. (1978) noted
a 5-day difference in mean gestation length between Scot-
tish red deer hinds managed under differing nutritional
regimens. More recently, Asher et al. (2005a) observed
marked variation in gestation length between hinds that
were fed either greater or lesser amounts of nutrients
during the third trimester of pregnancy. In that study ges-
tation length was negatively correlated with change in hind
live weight and the authors considered that under condi-
tions of modest feed imbalance that appeared to influence
foetal growth trajectory, a gestation length compensatory
mechanism ensured optimal birth weight. In effect, the
hypothesis argued that gestation length matched foetal
growth rate to ensure optimum birth weight (possibly at
the expense of optimal birth date).
Since the study of Asher et al. (2005a), other research has
demonstrated a statistically robust association between
conception date and gestation length for red deer sub-
species. Garcia et al. (2006) observed that Spanish red
174 G.W. Asher / Animal Reproduction Science 124 (2011) 170–175
deer hinds (C.e. hispanicus) that were artificially induced
to conceive up to 68 days before the standard mating date
exhibited longer gestation lengths than those conceiving
later in the natural breeding season. They postulated that
hinds were able to modify gestation length to counteract
out-of-season breeding to optimise calving date. However,
these previous data were confounded by the fact that early
conceiving hinds exhibited lower live weight gains over
pregnancy, thereby adding some support to the hypoth-
esis of Asher et al. (2005a) that nutrition was a driver of
gestation length variation. Most recently, Scott et al. (2008)
investigated the relationship between conception date and
gestation length in Scottish and Eastern European sub-
species of red deer (C.e. scoticus and C.e. hippelaphus), using
data for 393 naturally mated hinds across two herds, as well
as 91 hinds in which oestrus/conception were artificially
synchronised across a 4-week range of dates spanning the
natural rut. In all cases there were highly significant neg-
ative slopes for the regression of gestation length against
conception date. Effects for hind age, hind liveweight, calf
gender, birth weight, sire genotype and year were not sig-
nificant. The data supported a hypothesis that within-herd
synchrony of red deer births is facilitated by a ‘push/pull’
control over gestation length, such that hinds conceiv-
ing early and late in the breeding season have longer and
shorter gestation periods, respectively. The individual data
sets indicate that for every 10 days difference in concep-
tion date there was a change in gestation length of 1.9–4.9
days. The precise mechanisms remain to be resolved and
current studies by Scott et al. (unpublished data) are testing
the hypothesis that variation in fetal age during the latter
stages of pregnancy, when feed quality and voluntary feed
intake cycles of dams are in a state of flux, drives differential
growth trajectories for early and late conceiving foetuses,
leading to nutritional control over foetal maturation and
induction of parturition.
The aforementioned studies on red deer subspecies
indicate a hitherto unexpected environmental control over
gestation length in, at least some, cervids. Recent informa-
tion indicates that this may also be the case for reindeer
(Rowell and Shipka, 2009) and sika deer (Cervus nippon)
(Matsuura et al., 2004).
4. Conclusion
Clearly, there is no one ‘model’ species that defines typ-
ical cervid reproductive cycles, attesting to the complex
array of life histories and reproductive strategies of the
cervids in general. Our current understanding of reproduc-
tive biology largely centres around large-bodied temperate
(and highly seasonal) species that have a close association
with man. In contrast, reproductive cycles of the smaller-
bodied tropical (aseasonal) species are generally poorly
understood, with this group represented by many taxa that
are at serious risk of extinction.
Conflict of interest statement
The author has no conflict of interest in publishing this
review.
References
Asher, G.W., 1985. Oestrous cycle and breeding season of farmed fallow
deer, Dama dama. J. Reprod. Fertil. 75, 521–529.
Asher, G.W., 2007. Gestation length in red deer: genetically determined or
environmentally controlled? In: Juengel, J.L., Murray, J.F., Smith, M.F.
(Eds.), Reproduction in Domestic Ruminants VI. Nottingham Univer-
sity Press, Nottingham, UK, pp. 255–260.
Asher, G.W., Peterson, A.J., Watkins, W.B., 1988a. Hormonal changes dur-
ing luteal regression in farmed fallow deer, Dama dama. J. Reprod.
Fertil. 84, 379–386.
Asher, G.W., Adam, J.L., Otway, W., Bowmar, P., van Reenan, G., Mack-
intosh, C.G., Dratch, P., 1988b. Hybridisation of Pere David’s deer
(Elaphurus davidianus) and red deer (Cervus elaphus) by artificial
insemination. J. Zool. (London) 215, 197–203.
Asher, G.W., Fisher, M.W., Fennessy, P.F., Suttie, J.M., Webster, J.R., 1993a.
Manipulation of reproductive seasonality of farmed red deer (Cervus
elaphus) and fallow deer (Dama dama) by strategic administration of
exogenous melatonin. Anim. Reprod. Sci. 33, 267–287.
Asher, G.W., Fisher, M.W., Fennessy, P.F., Mackintosh, C.G., Jabbour, H.N.,
Morrow, C.J., 1993b. Oestrous synchronization, semen collection and
artificial insemination of farmed red deer (Cervus elaphus) and fallow
deer (Dama dama). Anim. Reprod. Sci. 33, 241–265.
Asher, G.W., Fisher, M.W., Berg, D.K., Waldrup, K.A., Pearse, A.J., 1996.
Luteal support of pregnancy in red deer (Cervus elaphus): effect of
cloprostenol, ovariectomy and lutectomy on the viability of the post-
implantation embryo. Anim. Reprod. Sci. 41, 141–151.
Asher, G.W., Muir, P.D., Semiadi, G., O’Neill, K.T., Scott, I.C., Barry, T.N.,
1997a. Seasonal patterns of luteal cyclicity in young red deer (Cervus
elaphus) and sambar deer (Cervus unicolor). Reprod. Fertil. Dev. 9,
587–596.
Asher, G.W., Scott, I.C., O’Neill, K.T., Smith, J.F., Inskeep, E.K., Townsend,
E.C., 1997b. Ultrasonographic monitoring of antral follicle develop-
ment in red deer (Cervus elaphus). J. Reprod. Fertil. 111, 91–99.
Asher, G.W., Monfort, S.L., Wemmer, C., 1999a. Comparative reproductive
function in cervids: implications for management of farm and zoo
populations. J. Reprod. Fertil. 54, 143–156.
Asher, G.W., Scott, I.C., Mockett, B.G., O’Neill, K.T., Diverio, S., Inskeep,
E.K., Townsend, E.C., 1999b. Ultrasonographic monitoring of ovarian
function during the oestrous cycle of fallow deer (Dama dama). In:
Proceedings of the Fourth International Congress on the Biology of
Deer, Kaposvar, Hungary, pp. 148–152.
Asher, G.W., Pearse, A.J., 2002. Managing reproductive performance of
farmed deer: the key to productivity. In: Proceedings of the Third
World Deer Farming Congress, Austin, TX, USA, pp. 99–112.
Asher, G.W., Mulley, R.C., O’Neill, K.T., Scott, I.C., Jopson, N.B., Littlejohn,
R.P., 2005a. Influence of level of nutrition during late pregnancy on
reproductive productivity of red deer (1) adult and primiparous hinds
gestating red deer calves. Anim. Reprod. Sci. 86, 261–283.
Asher, G.W., Scott, I.C., O’Neill, K.T., Littlejohn, R.P., 2005b. Influence of
nutrition during late pregnancy on reproductive productivity of red
deer (2) adult hinds gestating wapiti × red deer crossbred calves.
Anim. Reprod. Sci. 86, 285–296.
Berg, D.K., Thompson, J.G., Peterson, A.J., Asher, G.W., 1994. Morphol-
ogy and chronology of red deer (Cervus elaphus) preimplantation
embryonic development. Recent developments in deer biology. In:
Proceedings of the Third International Congress on the Biology of Deer,
Edinburgh, UK, pp. 179–180.
Berg, D.K., Thompson, J.G., Peterson, A.J., Asher, G.W., 2008. The temporal
relationship between oocyte maturation and early fertilization events
in relation to the pre-ovulatory LH peak and pre-implantation embryo
development in red deer (Cervus elaphus). Anim. Reprod. Sci. 105,
332–343.
Blouch, R.A., 1987. Reproductive seasonality of the white-tailed deer on
the Columbian Llanos. In: Wemmer, C.M. (Ed.), Biology and Manage-
ment of the Cervidae. Smithsonian Institution Press, Washington, pp.
339–343.
Chapman, D.I., Chapman, N.G., Dansie, O., 1984. The periods of concep-
tion and parturition in feral Reeves’ muntjac (Muntiacus reevesi) in
southern England based upon age of juvenile animals. J. Zool. 204,
575–578.
Chapple, R.S., English, A.W., Mulley, R.C., 1993. Characteristics of the
oestrous cycle and duration of gestation in chital hinds (Axis axis).
J. Reprod. Fertil. 98, 23–26.
Curlewis, J.D., Loudon, A.S.I., Coleman, A.M.P., 1988. Oestrous cycles and
the breeding season of the Père David’s deer hind (Elaphurus davidi-
anus). J. Reprod. Fertil. 82, 119–126.
Garcia, A.J., Landete-Castillejos, T., Carrion, D., Gaspar-Lopez, E., Gal-
lego, L., 2006. Compensatory extension of gestation length with
 G.W. Asher / Animal Reproduction Science 124 (2011) 170–175
advance of conception in red deer (Cervus elaphus). J. Exp. Zool. 305A,
55–61.
Guinness, F.E., Lincoln, G.A., Short, R.V., 1971. The reproductive cycle of
the female red deer, Cervus elaphus. J. Reprod. Fertil. 27, 427–438.
Guinness, F.E., Gibson, R.M., Clutton-Brock, T.H., 1978. Calving times of red
deer (Cervus elaphus) on Rhum. J. Zool. (London) 185, 105–114.
Haigh, J.C., 2001. The gestation length of wapiti (Cervus elaphus) revisited.
Anim. Reprod. Sci. 65, 89–93.
Harper, J.L., Clatworthy, J.N., McNaughton, I.H., Sagar, G.R., 1961. The evo-
lution and ecology of closely related species living in the same area.
Evolution 15, 209–227.
Harrington, R., 1985. Evolution and distribution of the Cervidae. In:
Fennessy, P.F., Drew, K.R. (Eds.), Biology of Deer Production. The Royal
Society of New Zealand, Bulletin 22, Wellington, New Zealand, pp.
3–11.
Kenneth, J.H., Ritchie, G.R., 1953. Gestation periods, a table and bibliog-
raphy. Technical Communications of the Commonwealth Bureau of
Animal Breeding and Genetics no. 5.
Lincoln, G.A., 1985. Seasonal breeding in deer. In: Fennessy, P.F., Drew, K.R.
(Eds.), The Biology of Deer Production. Bulletin No. 22, Royal Society
of New Zealand, Wellington, pp. 165–179.
Lincoln, G.A., Short, R.V., 1980. Seasonal breeding: nature’s contraceptive.
Recent Prog. Horm. Res. 36, 1–52.
Loudon, A.S.I., Curlewis, J.D., 1988. Cycles of antler and testicular growth
in an aseasonal tropical deer (Axis axis). J. Reprod. Fertil. 83, 729–738.
McCorkell, R.B., Woodbury, M.R., Adams, G.P., 2006. Ovarian follicular and
luteal dynamics in wapiti during the estrous cycle. Theriogenology 65,
40–56.
McMahon, C.D., Fisher, M.W., Mockett, B.G., Littlejohn, R.P., 1997. Embryo
development and placentome formation during pregnancy in red
deer. Reprod. Fertil. Dev. 9, 723–730.
Marshall, F.H.A., 1937. On the change over in the oestrous cycle in animals
after transference across the equator, with further observations on the
incidence of the breeding seasons and the factors controlling sexual
periodicity. Proc. Royal Soc. Lond. Ser. B 122, 413–428.
Matsuura, Y., Sato, K., Suzuki, M., Ohtaishi, N., 2004. The effects of age, body
weight and reproductive status on conception dates and gestation
periods in captive sika deer. Mamm. Study 29, 15–20.
Mishra, H.R., Wemmer, C., 1987. The comparative breeding ecology of four
cervids in Royal Chitwan National Park. In: Wemmer, C.M. (Ed.), Biol-
ogy and Management of the Cervidae. Smithsonian Institution Press,
Washington, USA, pp. 259–271.
Monfort, S.L., Wemmer, C., Kepler, T.H., Bush, M., Brown, J.L., Wildt, D.E.,
1991. Monitoring ovarian function and pregnancy in Eld’s deer (Cervus
175
eldi thamin) by evaluating urinary steroid metabolite excretion. J.
Reprod. Fertil. 88, 271–281.
Morrow, C.J., Penfold, L.M., Wolfe, B.A., 2009. Artificial insemination in
deer and non-domestic bovids. Theriogenology 71, 149–165.
Mylrea, G.E., 1992. Natural and artificial breeding of farmed chital deer
(Axis axis) in Australia PhD thesis, University of Sydney, NSW, Aus-
tralia.
Plotka, E.D., Seal, U.S., Verme, L.J., Ozoga, J.J., 1982. Reproductive steroids
in white-tailed deer IV Origin of progesterone during pregnancy. Biol.
Reprod. 26, 258–262.
Racey, P.A., 1981. Environmental factors affecting the length of gestation
in mammals. In: Cook, B., Gilmore, D.F. (Eds.), Environmental Factors
in Mammal Reproduction. Macmillan, London, pp. 199–213.
Rowell, J.E., Shipka, M.P., 2009. Variation in gestation length among
captive reindeer (Rangifer tarandus tarandus). Theriogenology 72,
190–197.
Schwartz, C.C., Hundertmark, K.J., 1993. Reproductive characteristics of
Alaskan moose. J. Wild. Manag. 57, 454–468.
Scott, I.C., Asher, G.W., Lach, J.E., Littlejohn, R.P., 2006. The influence of red
deer genotype on conception pattern: Eastern vs. Western subspecies.
Proc. New Zeal. Soc. Anim. Prod. 66, 270–273.
Scott, I.C., Asher, G.W., Archer, J.A., Littlejohn, R.P., 2008. The effect of con-
ception date on gestation length of red deer (Cervus elaphus). Anim.
Reprod. Sci. 109, 206–217.
Short, R.V., Hay, M.F., 1966. Delayed implantation in the roe deer, Capreolus
capreolus. Symp. Zool. Soc. Lond. 15, 173–194.
Sjaastad, O.V., Blom, A.K., Austad, R., Oen, E.O., 1990. Plasma progesterone
in reindeer in relation to ovariectomy and hysterectomy. Act. Vet.
Scand. 31, 45–51.
Smith, T.R., Hunter, C.G., Eisenberg, J.F., Sunquist, M.E., 1996. Ecology
of white-tailed deer in eastern Everglades National Park. Florida
Museum Natl. History 39, 141–172.
Van Mourik, S.A., Stelmasiak, T., 1990. Endocrine mechanisms and antler
cycles in rusa deer, Cervus (Rusa) timorensis. In: Bubenik, G.A., Bubenik,
A.B. (Eds.), Horns, Pronghorns and Antlers. Springer–Verlag, New York,
pp. 265–297.
Wemmer, C., Halverson, T., Rodden, M., Portillo, T., 1989. The reproductive
biology of female Père David’s deer (Elaphurus davidianus). Zoo Biol.
8, 49–55.
Whitehead, G.K., 1993. The Whitehead Encyclopaedia of Deer. Swan Hill
Press, Shrewsbury, UK.
Wong, B., Parker, K.L., 1993. Estrus in black-tailed deer. J. Mamm. 69,
168–171.