Behavioural factors affecting male reproductive success in red deer (Cervus elaphus)

Variation in reproductive success among adult red deer stags was related to harem size, and the duration and timing of breeding access. All three factors were affected by dominance (fighting success) during the rut. The number of hinds using a stag's rutting area and the duration of individual rutting activity also affected breeding access. Dominance rank in bachelor herds, which may affect access to hinds conceiving after the rut, was correlated with rank in the preceding but not the following rut.     

雄性麋鹿的交配机会、社会等级和皮质醇水平

我们在北京麋鹿苑研究了发情期雄性麋鹿的社会行为、社会等级和血清皮质醇水平,并分析了三者间的相互关系。按照雄性麋鹿的行为,将雄性麋鹿分为优势群和从属群。优势群包括群主和挑战者,从属群包括单身汉和被淘汰群主。使用SJ—Ⅰ型事件记录器进行常规行为取样,用放射免疫分析法测定血清皮质醇含量。发现：(1)优势雄鹿的发情行为频次明显高于从属雄鹿;(2)攻击行为的发生频次在两组间无显著差异;(3)血清皮质醇含量在两组间也无显著差异。由此我们认为,发情期雄性麋鹿的社会等级序位影响许多行为的表达,优势雄鹿成功交配的机会比从属雄鹿多。而攻击行为和肾上腺皮质醇的分泌与社群等级无明显相关,可能是麋鹿在圈养状态下的一种适应策略,这将有利于缓解发情期雄鹿的紧张,从而允许更多的能量投放在发情交配上。     

Relationship between Serum Testosterone, Dominance and Mating Success in Père David's Deer Stags

We conducted an experiment in the Beijing Milu Park to study the social behavior of male Père David's deer, and related social behavior to social position and serum testosterone level of the stags during rut. We classified the stags into three rank classes according to their rutting behavior: ‘harem master’, ‘challenger’ and ‘bachelor’. We monitored the behaviors of four ‘harem masters’, five ‘challengers’ and five ‘bachelors’, and analyzed serum testosterone levels in blood samples of those 14 stags using radioimmunoassay. We defined the effectiveness value, E = A/T, to assess the effectiveness of herding or mating attempts made by stags (‘T’ represents the frequency of herding or mating attempts made by a stag and ‘A’ represents the frequency of herding or mating attempts accepted by hinds). We found that: (1) the ‘harem masters’ and the ‘challengers’ displayed more frequent rut and locomotive behaviors but fewer ingestion behaviors than the ‘bachelors’; (2) serum testosterone levels in the ‘harem masters’ and the ‘challengers’ were higher than that in the ‘bachelors’; (3) effectiveness value of herding attempts differed significantly between the three types of stags, being highest in the ‘harem masters’ and the lowest in the ‘bachelors’; and (4) effectiveness value of mating attempts was significantly greater for the ‘harem masters’ than for the ‘challengers’. We conclude that: (1) reproductive behavior of the Père David's deer stags is strongly associated with social rank; (2) social roles of Père David's deer stags during the rut are related to the testosterone secretion; and (3) rank class affects the mating opportunity of the stags.     

Social dominance in farmed red deer stags

The dominance hierarchy was observed and recorded in a group of five farmed rising 5-year-old red deer stags, kept apart from hinds, from February 1982 until September 1982. The stags were observed at approximately fortnightly intervals during this period. The frequency of roaring and of serious fighting was also recorded. The identity of the dominant stag altered during the study on five occasions; a triangular dominance relationship was observed in only one observation period, and otherwise the hierarchy was linear. The roaring rate was high during the period when sexual activity would have taken place if hinds had been available, and the frequency of serious fighting was highest when one particular stag was dominant. Dominant stags showed no tendency to selectively harass subordinates in particular hierarchial positions. Although body weight was not related to rank at the beginning or end of the study, it was after 8 weeks had elapsed. Deer farmers who must keep adult stags together during the winter should maximise spacing to reduce aggression, and if necessary cull out trouble-makers.     

Red Deer (Cervus elaphus) Hinds Discriminate Between the Roars of Their Current Harem-Holder Stag and Those of Neighbouring Stags

During the period of reproduction red deer stags gather harems and roar loudly and repeatedly, both toward competing stags and toward the hinds that they actively herd. It has been proposed that red deer hinds may actively choose their mate on the basis of a comparison of the roaring rate of competing stags. Hinds may also choose to mate with the male that is most familiar to them, i.e. that male who spent most time and effort in retaining them within his harem and in roaring at them. Such a mechanism of female choice implies that females are able to discriminate individual characteristics in the stags' roars. We investigated this possibility by conducting playback experiments based on the habituation–discrimination paradigm. Our results show that hinds are able to discriminate between the roars of their current harem-holder stag and those of other neighbouring stags and suggest that this ability, a necessity for individual recognition, may be important in female mate choice in red deer.     

The induction of male sexual behavior in red deer (Cervus elaphus) by the administration of testosterone to hinds and estradiol-17β to stags

One-gram implants of testosterone were placed subcutaneously in two adult intact and one castrated red deer hinds, and 100-mg implants of estradiol-17β were given to three castrated and two intact red deer stags. Their sexual behavior was then observed in the wild and in captivity for up to 3 years. Testosterone initially produced prolonged and intense estrous behavior in the hinds; this gradually gave way to behavior normally shown only by rutting stags, such as flehmen, herding threats, and roaring. Testosterone-implanted hinds rose in the dominance hierarchy as measured by competition for food. Pregnancy, or the administration of progestagens, suppressed most of this testosterone-induced behavior. Testosterone also induced development of male secondary sexual characteristics such as formation of small antler pedicles (but not antlers), hypertrophy of the clitoris and neck musculature, and development of a neck mane and male rutting odor. Estradiol-17β mimicked the behavioral effects of testosterone when given to castrated stags, stimulating all components of rutting behavior. In intact stags, the only effects were to abolish antler casting and stimulate roaring; normal rutting behavior continued unchanged. In contrast to the effects of testosterone, estradiol-17β did not influence the social status of either intact or castrated stags. The distinctions between the behavioral effects of androgens and estrogens are ill defined and are determined more by the prior sexual differentiation of the brain and the duration of steroid treatment, than by the nature of the steroid.     

Induction of early breeding in red deer (Cervus elaphus) by melatonin

Lactating (N = 12) and non-lactating (N = 6) red deer hinds and one stag at pasture were given concentrates (500 g/head) containing melatonin (5 mg/head) daily at 16:00 h from 18 June to 16 October. The stag shed the antler velvet and started rutting 5 weeks ahead of untreated stags, and hinds had their first oestrus and ovulation of the breeding season in mid-September, 5 weeks in advance of control lactating (N = 9) and non-lactating (N = 5) hinds. Treated hinds were allowed to mate with the treated stag and control hinds ran in an adjacent paddock with an untreated stag. All hinds became pregnant during the study with all but 2 melatonin-fed hinds (1 lactating, 1 non-lactating) and 1 control hind (lactating) conceiving at the first oestrus. Melatonin-treated hinds lost slightly more live weight than did controls from June to November, but their suckled calves grew throughout at a rate similar to those of control hinds.     

"占群"还是"挑战"?不同时间限制条件下麋鹿个体的交配计策

在麋鹿的发情交配季节 ,雄性麋鹿可区分为 3种类型 :“群主”、“挑战者”和“单身汉”。“群主”是一头圈占并控制雌鹿活动的优势雄性。“挑战者”不占有雌性繁殖群 ,但在发情场附近地点展示炫耀。当雌性繁殖群的雌鹿外出采食靠近“挑战者”的展示炫耀地点时 ,“挑战者”会积极地寻求机会与之交配。“单身汉”在繁殖季节不表现发情行为。他们像非繁殖季节一样采食 ,采食后蹲在水塘中休息。我们对何种因素决定麋鹿个体的发情交配计策感兴趣。 1996至 1998年夏天 ,我们在北京麋鹿苑观察麋鹿发情交配行为以分析导致这些行为差异的原因。结果发现 ,“群主”、“挑战者”和“单身汉”用于维持生命的时间预算与用于发情的时间预算成反比 ,并且 ,“群主”、“挑战者”和“单身汉”用于维持生命的时间预算与用于发情的时间预算差异显著。“群主”的绝大部分时间用于发情占群 ,而用于采食、饮水的时间很少 ,所以 ,“群主”在发情期间基本上处于禁食状态 ,靠消耗体内脂肪维持生命。“单身汉”则相反 ,绝大部分时间用于采食、休息和反刍 ,基本上没有发情行为。“挑战者”在发情行为与维持生命行为之间的时间则居于“群主”与“单身汉”之间。交配次数是偏态分布的 ,与雄性发情时间呈正比。“群主”的交配概率最高     

Expression and relevance of dominance in farmed Rusa deer (Cervus rusa timorensis)

A group of 50 hinds and four Rusa stags (Cervus rusa timorensis) were observed over a 2-year period under farming conditions. The aim of the study was to investigate the expression and relevance of dominance. A dominance order was calculated for the hinds. No significant influence of dominance value in relation to calving time, presence or absence of calf, mating time for the following season or the age of the hinds was found. The number of victories in comparison to the total number of interactions between stags was influenced by possession of a full set of antlers for young as well as old stags. However, the antler size or shape did not affect successful participation in the rutting activities of the stags. Hinds did not show a preference for stags based on antler appearance. It is recommended that stags should be velveted so as to minimize differences in antler shape and consequently to reduce dominance interactions between them.     

Artificial insemination following observational versus electronic methods of estrus detection in red deer hinds (Cervus elephus)

The objectives were to determine the efficacy of the HeatWatch (HW) electronic estrus detection system for monitoring behavioral estrus (including duration and intensity) in red deer hinds and to evaluate pregnancy rate to AI after detected estrus. Red deer hinds (Cervus elephus; n = 50) were allocated into two treatment groups: AI following synchronization (CIDR/PMSG) and observed estrus (induced estrus group: IE; n = 25) or AI following the detection of natural estrus (NE; n = 25) without hormonal treatment. Hinds were fitted with two HeatWatch (HW) electronic estrus detection transmitters, one above the tail (bottom) and one between the tuber coxae of the pelvic girdle (top), and visual observations for mounting activity began with the aid of young sterile red deer stags (18 months old) fitted with marking harnesses. Hinds in both groups were inseminated (10-12h after observed estrus) with frozen-thawed red deer semen using a transvaginal/cervical AI approach. Following a 26-day period of AI, hinds were placed with a mature fertile stag for an additional 30-day natural breeding period. Pregnancy diagnosis was performed 57 and 86 days after the start of AI. While the hinds were housed with the young stags, 82% were detected in estrus by visual appraisal of stag crayon marks, but only 32% of these were detected by HW. In contrast, in the hinds housed with the mature stag, 93% detected in estrus by crayon marks were also detected by HW. The top HW transmitter consistently recorded more mounts (P < 0.05) than the bottom transmitter. The pregnancy rate was numerically better in IE versus NE hinds (42% versus 29%, P > 0.10). In summary, there were no differences (P > 0.10) in the intensity (number) or duration of mounts (detected by HW) during estrus in IE versus NE hinds, and HW was most effective in detecting estrus in the presence of a heavier, mature stag versus a younger stag. When used in combination with transvaginal AI, an overall first-service pregnancy rate of 36.6% was achieved with AI of frozen-thawed semen in red deer hinds following detected estrus.     

The effect of cyproterone acetate on the antler cycle in red deer (Cervus elaphus L.)

The aim of the study was to test the effect of antiandrogen, cyproterone acetate (CA) on the antler cycle in the red deer (Cervus elaphus). CA was administered to three adult red deer stags (Edward, Fuks and Gacek) in weekly intervals. Edward and Fuks were given 600 mg + 600 mg of CA, whereas Gacek was given 600 mg + 300 mg. CA was injected during the hard antler phase: in mid-October (Edward), at the end of November (Fuks) and at the end of January (Gacek). CA caused the antler casting 17 to 22 days after the first injection. In all stags, the casting of antlers was followed by a period of intensive growth of new antlers. Edward was given CA at the end of October. This treatment was responsible for occurrence of the two antler cycles in the year of the experiment. When CA was administered during the middle of the hard antler phase an additional short antler cycle occurs followed by new antler growth. CA treatment in the later part of hard antler phase may cause a prolonged antler cycle.     

Induction of antler growth in a congenitally polled Scottish red deer stag.

A congenitally polled red deer stag was captured from a Scottish deer forest and kept in an enclosure for observations. The animal had rudimentary antler pedicles but no antlers, and during five years of study no significant antler development occurred. Amputation of the apex of one antler pedicle in May 1974 when the stat was 12 years of age resulted in the growth of a complete antler on the operated side, and this antler was subsequently cleaned and cast in the normal way and a new antler cycle was initiated. The result illustrates that the primary abnormality in this polled stag lay not in his inability to grow antler, but in his inability to develop fully formed antler pedicles from which normal antler tissue could differentiate. Traumatizing the rudimentary pedicle had the effect of stimulating growth of antler tissue, and once this was formed the process of cleaning, casting and regrowth occurred spontaneously. The incomplete development of the antler pedicles is considered to be responsible for the absence of antlers in the majority of "hummels" in Scotland, and the etiology of the condition is discussed.     

Effects of melatonin implants on reproductive seasonality of male red deer (Cervus elaphus)

Red deer stags were treated with melatonin implants in 2 experiments designed to examine the control of reproductive seasonality. In Exp. 1, stags (n = 24) were allocated to 4 treatment groups: 2 groups were treated with 3 implants per stag each month from 8 November to 5 February (EM) or 9 December to 5 February (LM), 1 untreated group of control stags remained with the melatonin-treated stags (CC) and the other untreated control group remained isolated (IC). Melatonin treatment advanced the seasonal changes in scrotal circumference, liveweight, antler state and coat type compared with control stags. The extent of advancement was greater in EM than LM stags. In EM and LM stags, size of testes regressed rapidly and antlers were cast shortly after melatonin implants became exhausted in March. This was followed by an additional antler cycle and reproductive development and decline from June to November. EM and LM stags became synchronized with control stags 14-15 months after melatonin treatment began. The extra cycle of seasonal changes was more pronounced in EM than in LM stags. In Exp. 2, stags (n = 30) were allocated to 6 treatment groups: 4 groups were treated with 3 implants per stag at monthly intervals for 6 months from 22 June (J), 4 August (A), 16 September (S) and 23 October (O), a further group of stags was treated in the same manner for 12 months from 22 June (Y), and the remaining group was untreated (C). Compared with control stags, testicular regression and antler casting was delayed in Groups J, A and Y. These events occurred at the same time as in control stags in Groups S and O. Subsequent reproductive development was advanced in Groups S and O and delayed in Groups J, A and Y. The results demonstrated that treatment with melatonin implants in November or December advanced reproductive development. However, when stags were treated with melatonin implants from June to August, reproductive development was delayed, indicating a change in response to melatonin treatment during the year. The change in response to melatonin treatment between late winter and early spring was interpreted as a resetting of an endogenous circannual rhythm caused by a photoperiodic cue responsible for initiating the final stages of reproductive regression.     

Antler size and fluctuating asymmetry in red deer (Cervus elaphus) stags and probability of becoming a harem holder in rut

Red deer hinds play a significant role in selecting stags for mating. We tested the hypotheses that in red deer ( Cervus elaphus ) the probability of becoming a harem holder (and hence achieving reproductive success) occurs in stags bearing larger, branched antlers and showing low fluctuating asymmetry (FA). Eleven antler characteristics were measured; absolute and relative FA were calculated on 51 cast antler sets from 19 individually recognized stags. Probability of becoming a harem holder (PBHH) was originally analysed separately, i.e. for antler size and FA of each antler characteristic and calculated factors for both antler size and FA. If analysed separately, large antler size and low relative but high absolute FA increased PBHH. When we combined antler size and FA of antler characteristics in one model using antler size and FA factors, however, PBHH and achieving reproductive success were mainly dependent on increasing antler size and enhanced antler branching rather than on FA. We conclude that in contrast to antler size, FA is unlikely to play any significant role in sexual selection as an indicator of individual quality.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 59–68.     

Great expectations: dominance,breeding success and offspring sex ratios in red deer

Differences in dominance rank among red deer (Cervus elaphus) females (hinds) on Rhum were related to their breeding success as well as to the comparative success of male and female offspring. Males (stags) born to mothers above median rank were more successful than hinds, while hinds born to subordinate mothers were more successful than stags. The ratio of male to female calves produced by dominant mothers was significantly higher than that produced by subordinates. Since dominance rank among hinds is related to their body weight as adults and to their birth weight, these results suggest that the birth sex ratio may be affected by environmental factors operating during a female's early development.     

Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland

A pattern of wolfCanis lupus Linnaeus, 1758 predation on red deerCervus elaphus Linnaeus, 1758 was studied in Bieszczady Mountains in 1991–2002. In total 324 remains of red deer > 4 months old, killed by wolves throughout the year, were found. The sex, age and bone marrow fat content of wolf kills were compared with the same characteristics within the free living red deer population. The overall contribution of calves killed by wolves (24%) in October-May was higher than in the population (17%), and decreased from autumn to spring. Adult males were more vulnerable to wolf predation than adult females: stags constituted 62% and hinds 38% of adult red deer killed by wolves, whereas in the population, the percentages were 37 and 63%, respectively. Stags killed by wolves were younger ([`(x)] = 4.1\bar x = 4.1 years old) than hinds ([`(x)] = 8.9\bar x = 8.9 years old). Wolves killed more > 8 years old hinds and < 5 years old stags than available in the population. In wolf kills, the average fat content in femur marrow was higher among hinds (84.9%) than stags (69.3%) and calves (66.1%). Only 8% of hinds had < 70% femur marrow fat content, whereas 40% of calves and 38% of stags had marrow fat values below that level. Marrow fat content showed seasonal variation and was the lowest in March among all sex-age classes. The monthly share of stags in all kills, and hinds in hind-calf part of the sample was negatively correlated with their monthly average bone marrow fat content, and monthly share of calves was positively correlated with monthly average bone marrow fat content of adults. The segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter until spring contribute to the high overall incidence of calves and adult males and the relatively low incidence of adult females among wolf kills.     

Seasonal changes in fecal testosterone concentrations and their relationship to the reproductive behavior, antler cycle and grouping patterns in free-ranging male Pampas deer (Ozotoceros bezoarticus bezoarticus)

The purpose of this study was to validate noninvasive endocrine monitoring techniques for Pampas deer and to evaluate seasonal changes in testicular steroidogenic activity and their correlation to reproductive behavior, antler cycle and group size. Thus, fecal samples, behavioral data and observations of antler status were collected at monthly intervals during 1 year from free-ranging Pampas deer stags (three radio-collared individuals and 15 random individuals) living in Emas National Park, Brazil (18 degrees S latitude). Fecal steroids were extracted using 80% methanol and steroid concentrations were quantified by a commercial enzyme immunoassay (EIA). Fecal testosterone concentrations peaked in December-January (summer), March (early autumn) and in August-September (winter-spring), with minimal values from April-July. Reproductive behavior had two peaks, the first in December-January, characterized by predominately anogenital sniffing, flehmen, urine sniffing, chasing and mounting behavior, and the second peak in July-September (behavior primarily related to gland marking). There were significant correlations between fecal testosterone and reproductive behavior (r=0.490), and between fecal testosterone and antler phases (r=0.239). Antler casting and regrowth occurred under low testosterone concentrations, whereas velvet shedding was associated with high concentrations of testosterone. We inferred that Pampas deer stags exhibited a seasonal cycle that modulated sexual behavior and the antler cycle, and we concluded that fecal steroid analysis was a practical and reliable non-invasive method for the evaluation of the endocrine status of free-ranging Pampas deer.     

Distribution and status of White-lipped Deer (Cervus albirostris) in the Qinghai-Xizang (Tibet) Plateau, China

We made a preliminary survey of the present status and ecology of the White-lipped Deer (Cervus albirostris) in the upper Huang He and Chang Jiang River regions of China. Eight mixed herds, ranging in size from 5 to 46 animals each, were observed. The deer lived from the rhododendron scrub just above the spruce forests to alpine grasslands, ranging in altitudes from 4000 to 5000 m. The peak of the rut occurred in October, when breeding herds were composed of a few large stags and hinds. The pasturing of domestic animals in the area influences not only the spatial distribution of the deer, but also the timing of seasonal shifts. The main distribution of the deer is in western Sichuan Province and some scattered populations in Qinghai Province are endangered. The species could survive, given adequate protection.     

Effects of antler breakage on mating behavior in male tule elk (<Emphasis Type="Italic">Cervus elaphus nannodes</Emphasis>)

Although antler size has been identified as a primary determinant of dominance, fighting success, and reproductive success in male cervids, >80% of the male tule elk (Cervus elaphus nannodes) in the Owens Valley, California, experience antler breakage. To determine the effect of antler breakage on male mating success, we recorded antler morphology, body size, and mating behavior of male elk throughout the rut. Antler breakage, regardless of severity, had no effect on male–male assessment, fighting success, or harem-holding status. The factor consistently associated with our indices of male mating success was not antler size but body size. Although antler size is frequently emphasized as a key factor in male dominance and social rank, this association may reflect the correlation between antler size and body size. In the Owens Valley, it appears that male elk are not assessing competitors based on antler morphology but on other characteristics. An erratum to this article can be found at     

Haematology and serum biochemistry of captive unsedated chital deer (Axis axis) in Australia

Haematological and serum biochemical values were measured in blood samples collected over a 12-mo period from 37 unsedated chital deer (Axis axis). Stags and hinds, ranging in age from birth to maturity, were sampled. Haemoglobin, total erythrocyte and haematocrit values were low at birth and sex differences were not apparent in neonates and juveniles, but were in adults. Chital stags had higher erythrocyte parameters (P less than 0.001) and lower erythrocyte indices than hinds, and the total leucocyte count was higher in stags (P less than 0.01). Some parameters (erythrocytes, muscle enzymes, glucose, cortisol) decreased over successive serial sampling. The differential leucocyte count of older stags decreased during the initial handling period. The major rutting period in February and March was characterised by changes in the differential leucocyte count, elevations in serum muscle enzymes, and lower serum cortisol levels. Alkaline phosphatase activity in serum reflected the annual antler cycle of chital stags. Serial sampling over many weeks, either weekly or tri-weekly, produced haematological and biochemical changes in successive samples which may have reflected a reduction in stress and excitement associated with restraint.