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.     

Relationships between Behaviour and Antler Cycle Timing in Red Deer

The factors affecting the stag social hierarchy and antler cycle timing in a herd of red deer were analysed. The analysis was based on behaviour records of the herd between 1972 and 1983. The alpha stag was not usually the oldest stag present. Changes in rank were commoner in periods of hard antler than during velvet periods. The frequency of rank changes was correlated with the number of stags present but not with the number of hinds; it was also positively related to indicators of the level of aggression such as the incidence of antler breakage, the number of harem-holding stags per rut and the incidence of post-casting rank loss. The number of stags killed in fights was not correlated to any of the indicators of aggression except the number of harem-holders per rut. Rank loss after antler casting and the change of harem-holding stags during the rut were both relatively rare events indicating year-round, stable social relationships in the herd. The relationships between rank and casting date and between rank and cleaning date previously reported were related to the level of aggression among the deer. The higher the level of aggression, the closer the relationship between rank and the timing of the antler cycle. The small difference in the level of aggression between the periods of casting and cleaning is thought to be the result of the seasonally different social background. The number of hinds present did not appear to affect antler casting, antler cleaning or the general indicators of the level of aggression. It is concluded that the results support the previously advanced hypothesis that behaviour modulates antler cycle timing in Cervids.     

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.     

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.     

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.     

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.     

Social Organization in an Enclosed Group of Red Deer (Cervus elaphus L.) on Rhum. I. The Dominance Hierarchy of Females and their Offspring

Studies of the dominance hierarchy in red deer have mainly concentrated on the hierarchy among stags. This study aimed to investigate in detail the dominance hierarchy of a group of hinds and their offspring and to compare it between two feeding situations, one when the deer were browsing normally in a large area, and the other when they were competing for artificially provided food in a small area.     

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.     

四川梅花鹿发情期的几种发声行为

2006 年9 ～11 月在四川省若尔盖县铁布自然保护区,对四川梅花鹿发情期的声行为进行了初步观测,结果表明:在发情期,四川梅花鹿的发声行为可分为雌、雄鹿的报警叫声,雄鹿的吼叫声和求偶叫声。雌鹿的报警叫声持续时间257 ～ 539 ms,频率范围1409. 5 ～ 4474. 6 Hz,主频率3534. 8 ±89.12 Hz;雄鹿的报警叫声持续时间136 ～187 ms,频率范围271.8 ～3910.5 Hz,主频率3244.3 ±79.32 Hz;两者在持续时间、最低频率、最高频率上差异极显著(P < 0.01),在间隔时间上差异不显著(P = 0.624)。吼叫是雄鹿的主要发情行为之一。雄鹿每次吼叫1 声,持续时间1580 ～4972 ms,频率范围234.6 ～6171.4 Hz,主频率2264.6 ± 166.44 Hz。雄鹿吼叫声的主频率存在显著的个体差异(P <0.01)。在整个吼叫过程中,一只雄鹿的吼叫常会引起周围其它雄鹿的吼叫反应。雄鹿每日吼叫的次数与其在繁殖群中的等级序列有关,不同序列等级雄鹿的吼叫频次存在显著差异(P<0.01)。雄鹿的吼叫声在白天和夜晚均能听到,但主要发生在06:00 ～ 08:00、17:00 ～ 19:00 和01:00 ～ 03:00 三个时间段。四川梅花鹿雄鹿的求偶叫声有4 种,其生物学意义与发情炫耀、追逐、激惹和爬跨等行为有关。     

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.     

Behavioral Dominance and Corpus Luteum Function in Red DeerCervus elaphus

Times of the ovulatory LH surge and characteristics of the rise in circulating progesterone concentrations after ovulation in red deer hinds were investigated in relation to each animal's dominance status. Observations were made during the 1992 (experiment 1) and 1993 (experiment 2) breeding seasons, while the same 12 hinds were held in a pen in the absence of a stag. Ovulation was synchronized by administration of progesterone followed by luteolytic prostaglandin F_2αanalogue. Social status was determined for each hind by noting dyadic agonistic interactions during the period of progesterone treatment. Hinds were weighed before and after the experiments. Time of onset of estrus (lordosis) was recorded while handling at 3-hr intervals (for 81 hr in experiment 1; 96 hr in experiment 2) after progesterone withdrawal, and blood samples were collected at these times to characterize the preovulatory LH surge. Subsequently, daily blood samples were collected for up to 11 days for measurement of progesterone concentrations. There was a tendency for weight change to be related to dominance status in experiment 1 and this was significant in experiment 2 (p< 0.01). The rate of increase in circulating progesterone concentration after ovulation was related to status (data of experiments 1 and 2 combined;p< 0.02), but was not correlated with the time of estrus or with the time or height of the LH surge. A third experiment (carried out in 1993), when the same hinds were kept with a stag after induced ovulation, showed that time of estrus (mating) was not related to dominance status. The data suggest that corpus luteum function is affected by social status. The results are discussed in the context of mechanisms by which dominance status influences the sex of a hind's calves.     

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.     

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。     

"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     

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.     

Reproductive performance of farmed red deer (Cervus elaphus) in New Zealand. I. Descriptive data.

A longitudinal observational study of 15 red deer (Cervus elaphus) farms was carried out in New Zealand for 2 years from Mar 1992. Management of hind mobs and their composition during mating, calving, and weaning were recorded. About 2700 hinds were individually monitored for live weight, body condition score and reproductive success. All hinds were pregnancy tested in June by ultrasound and classified as having conceived before May 1, after May 1, or as being not pregnant. Calving dates and dam-offspring pairs were recorded on four farms. The lactational status of hinds and live weight of calves were recorded at weaning. Mean pre-mating live weights of yearling hinds were 81.3-82.8 kg and of adult hinds 98.0-98.7 kg, respectively, for each year of study. The percentage of yearling and adult hinds conceiving before May ranged from 8.3 to 95.0% and from 77.6 to 98.4%, respectively, between farms. Overall, the proportion of yearling and adult hinds not pregnant was 15.3% and 3.2%, respectively. Pregnancy rates at scanning of adult hinds within mating mobs were generally over 90%, and 41.7% of mating mobs had 100% pregnancy rates. In contrast, the pregnancy rates of yearling hind mobs were more variable, with five mobs below 50%, and 34.1% of mating mobs achieving 100% pregnancy rate. Estimated in utero loss rates from pregnancy diagnosis to calving were 0.66 and 0.79% for yearling and adult hinds, respectively. From four farms, median calving dates of yearling and adult hinds were Dec 13 and Nov 30, respectively. Overall, calf survival rates to weaning of yearling and adult offspring were 84.1 and 91.6%, respectively, while the reproductive efficiencies (number of calves weaned per hind mated) were 70 and 83.6%, respectively. Farm mean weaner live weights standardised on Apr 1 ranged from 42-59 kg and 39-51 kg for stags and hinds, respectively. These data are currently the best estimates of reproductive parameters of New Zealand red deer herds, and highlight a wide variability in reproductive indices between farms.     

Assessment of dominance hierarchy through urine scent marking and its chemical constituents in male blackbuck Antelope cervicapra, a critically endangered species

In ungulates the process of chemical communication by urinary scent marking has been directly related to reproductive dominance, territorial defense and proximity to resources. The differences in the frequency of urine marking and chemical composition of urine of males Antelope cervicapra before, during and after the dominance hierarchy period were assessed. The variations in the urine marking and its chemical profiles of dominant males (n = 9), bachelors (n = 5) and sub-adult males (n = 5) were compared to find out how the dominance hierarchy influences the confined blackbuck herd under semi-natural captive conditions. The frequency of urine marking is significantly higher (p < 0.001) in dominant males. Twenty-eight major constituents were identified in the urine of dominant males (before, during and after the dominance hierarchy period), bachelor and sub-adult males. Among these, three specific compounds namely, 3-hexanone (I), 6-methyl-5-hepten-2-one (II) and 4-methyl-3-heptanone (III) were seen only in dominant males urine during the dominance hierarchy period. Based on the behavioural observation and the unique chemical constituents in the urine, it is concluded that the dominant male scent odor suppresses aggression, scent marking, scent production and territorial patrolling activities of subordinate males, through which the dominant male establish their hierarchy and attains success in reproduction.     

Oestrous cycles and the breeding season of the Père David's deer hind (Elaphurus davidianus)

Reproductive cycles were studied in a group of tame Père David's deer hinds. The non-pregnant hind is seasonally polyoestrous and, in animals studied over 2 years, the breeding season began in early August (2 August +/- 3.3 days; s.e.m., N = 9) and ended in mid-December (18 December +/- 5.7 days; N = 8) and early January (6 January +/- 3.2 days; N = 11) in consecutive years. During the anoestrous period, plasma progesterone concentrations were low (0.2 +/- 0.01 ng/ml) or non-detectable. There was a small, transient increase in progesterone values before the onset of the first cycle of the breeding season. In daily samples taken during an oestrous cycle in which hinds were mated by a marked vasectomized stag, progesterone concentrations remained low (less than 0.5 ng/ml) for a period of about 6 days around the time of oestrus, showed a significant increase above oestrous levels by Day 4 (Day 0 = day of oestrus) and then continued to increase for 18 +/- 2.8 days to reach mean maximum luteal levels of 3.5 +/- 0.6 ng/ml. The plasma progesterone profiles from a number of animals indicated that marking of the hinds by the vasectomized stag did not occur at each ovulation during the breeding season and therefore an estimate of the cycle length could not be determined by this method. In the following year, detection of oestrus in 5 hinds was based on behavioural observations made in the absence of the stag. A total of 19 oestrous cycles with a mean length of 19.5 +/- 0.6 days was observed.(ABSTRACT TRUNCATED AT 250 WORDS)     

Dominance hierarchy among workers changes with colony development in Polistes japonicus (Hymenoptera, Vespidae) paper wasp colonies with a small number of workers

Dominance hierarchy in the primitively eusocial wasp Polistes japonicus was analysed in four colonies for two periods: (1) the first-brood period, when only early emerging workers are present on the nest, and (2) the mixed-brood period, when the first and second (last) broods are present on the nest. The rank in the dominance hierarchy was determined based on a sociogram showing a dominance–subordinance relationship for all pairs of workers. During the first-brood period, older workers were likely to be more dominant (older dominance hierarchy), while the rank of workers was reversed during the mixed-brood period, with younger workers being likely to be more dominant (younger dominance hierarchy). However, the oldest and youngest workers were not always the top-ranked workers in the dominance hierarchy during the first- and mixed-brood periods, respectively, and during the mixed-brood period no younger dominance hierarchy was evident when the first or second brood was analysed separately. Higher ranked workers displayed dominance behaviour more frequently, and the lowest ranked worker hardly displayed dominance behaviour. Most workers displayed dominance behaviours primarily towards the worker ranked immediately below in the dominance hierarchy during the mixed-brood period but not during the first-brood period. The bodies of younger workers were larger for the mixed brood, but not for the first brood in some colonies or the second brood in all colonies. The association between body size and rank in the dominance hierarchy was negative during the first-brood period and positive during the mixed-brood period, with a nearly significant trend also seen even when the analysis was limited to the second brood. To explain the above temporal change from an older dominance hierarchy to a younger dominance hierarchy, we propose the hypothesis that the probability of a worker inheriting the colony increases rapidly with colony development, and consequently younger larger workers attempt to move up the dominance hierarchy in order to produce their own offspring by becoming the superseder late in colony development, rather than working harmoniously so as to boost the overall production of reproductive progeny for a colony, which is the strategy adopted early in colony development.