雄性麋鹿的吼叫行为、序位等级与成功繁殖

为探讨雄性麋鹿（Elaphurus davidianus）的吼叫行为与序位等级的关系,以及这种关系在成功繁殖中的作用。2000年7－8月,在北京麋鹿苑选定12头成体雄鹿,用目标取样法观察记录每头鹿的吼叫、圈群和交配行为的发生频次。用有效比值[被雌鹿接受的频次／雄鹿圈群（或交配）的总频次]来衡量雄鹿圈群行为（或交配行为）的有效性。依据雄鹿在发情期的表现状态,把雄鹿归为3类：占有雌鹿群的“群主”（4头）、未占有雌鹿群而挑战“群主”的“挑战者”（5头）和远离繁殖群的“单身汉”（3头）。不同序位等级雄鹿的吼叫频次、圈群（或交配）行为频次、圈群行为（或交配行为）的有效值存在显著差异（P＜0.05）。对吼叫行为的分析表明：“群主”吼叫频次最高,“单身汉”吼叫频次最低。“群主”的有效值最高,“单身汉”的有效值最低。据此认为雄性麋鹿的吼叫行为与其序位等级密切相关,而序位等级又决定了雄鹿参与繁殖的机会。此外,还记录到占群雄鹿的警戒吼叫,这是对Wemmer et al.(1983)警戒吼叫行为仅见于雌鹿的观察结果的补充。     

"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     

雄性海南坡鹿发情期泥浴、沙浴行为及其在繁殖中的作用

2005年2-5月间,在海南省大田国家级自然保护区文昌保护站的养鹿场内对7只半野生雄性海南坡鹿(Cervus eldi hainanus)的泥浴和沙浴行为、进行泥浴和沙浴行为的雄性个体的年龄、等级序位和交配成功次数进行了观察记录,以确定雄性坡鹿的泥、沙浴行为是否与雄性个体的年龄、等级序位和交配行为相关.本实验对以下3个假说进行了检验:(1)性成熟的雄鹿的泥浴行为多于刚刚性成熟的雄鹿;(2)优势雄鹿泥浴的频次和持续的时间多于非优势个体;(3)泥浴的次数与交配次数正相关.研究期间记录到雄性个体的79次泥浴和171次沙浴行为,性成熟雄鹿的泥浴和沙浴的频次显著地高于刚刚性成熟的雄鹿(P＜0.01,Friedman nonparametric two-way ANOVA);不同等级序位的雄性泥沙浴持续时间没有明显的差异,但是优势雄鹿的泥浴频次显著高于非优势个体(Wilcoxon signed ranks 检验,P＜0.01);雄鹿的泥浴和沙浴的频次分别与交配次数显著正相关(r=0.802, P＜0.05, n=7;r=0.919, P＜0.01, n=7;Pearson 检验).雄性海南坡鹿沙浴与泥浴是雄鹿发情期中的优势炫耀行为,对雄性个体的繁殖成功与否具有重要的意义[动物学报 53(3):417-424,2007].     

布氏田鼠雄性的优势地位

在实验条件下对雄性成年布氏田鼠（Microtus brandti）相遇后的优势地位及生理指标的研究结果表明：布氏田鼠雄性之间存在严格的等级关系;优势雄鼠生理指标中睾丸重量和血清睾酮激素含量高于从属雄鼠,而血清中皮质醇激素的含量低于从属雄鼠。优势雄鼠的探究、攻击和修饰等行为频次显著高于从属雄鼠,而从属雄鼠在跳跃频次上多于优势雄鼠。随着观察时间的延长,不论优势个体还是从属个体,其探究频次都呈现明显的下降趋势,其他行为的频次则没有显著差异。相关性检验说明,除攻击行为可以作为判断区分雄鼠等级的依据之外,修饰行为也可以作为判断的依据。     

不同密度下和不同发情期时段中雄性麋鹿的繁殖计策

从1998年到2001年,我们研究了密度、发情期与雄性麋鹿繁殖计策的关系。我们选择了江苏大丰麋鹿保护区的两个麋鹿群体和北京麋鹿苑的一个麋鹿群体,观察记录了发情期前期、中期和后期中实验麋鹿种群参数、围栏大小、群体中雄鹿的繁殖计策、群主变更情况、后宫数目以及后宫雌鹿数。尽管不同种群和不同发情期的雄鹿繁殖计策有差异,但是各个群体中雄性麋鹿都有占群、偷袭交配和求偶集会3种繁殖计策。利用因子分析提取了两个主成分:主成分1主要代表实验围栏、麋鹿存栏数和密度等因子的影响,主成分2主要代表发情期时段因子的影响。这两个主成分解释了全部因素效应的90.72%。用Spearman相关分析发现,占群计策与主成分2呈极显著负相关,求偶集会计策与主成分1呈显著正相关,最大后宫雌鹿个体数与主成分1极显著正相关,而偷袭交配计策和后宫数目与主成分1和2均无显著相关。据此推测,占群和求偶集会繁殖计策分别受密度和发情期时段的影响,而偷袭交配行为对密度和发情期时段不敏感。我们还探讨了麋鹿发情行为计策的环境适应性,从行为学角度为麋鹿保护提供了新思路。     

圈养雄性麋鹿血清睾酮和皮质醇含量在发情期的变化

2000年7-8月在北京麋鹿苑采集不同繁殖状态雄鹿（发情前3头、发情3头和发情后4头）血液样品,用放射免疫分析法测定血清睾酮和皮质醇含量。结果显示发情雄鹿的睾酮浓度显著高于发情前、后,而3种状态雄鹿的皮质醇浓度无差异,并且与睾酮浓度无关。本苑麋鹿睾酮水平的变化与同类野生动物一致。     

不同社群条件下雄性布氏田鼠的行为

实验室内观察了布氏田鼠雄性个体之间在3 种不同社群条件下(社群1:2 只雄鼠;社群2:2 只雄鼠与1只非动情雌鼠,社群3:2 只雄鼠与1 只动情雌鼠)的相互行为,结果显示:(1)在嗅闻行为上,不论优势雄鼠还是从属雄鼠在3 组间都存在显著差异,它们的嗅闻频次都是在社群3 中最少、在社群2中居中,在社群1中嗅闻频次最多;(2)在攻击行为频次上,不论是优势雄鼠还是从属雄鼠在3 组中都不存在显著差异;(3)在上跳频次上,优势雄鼠在各组间存在显著差异,从属雄鼠在各组间不存在显著差异; (4) 在自我修饰行为频次上,不论是优势雄鼠还是从属雄鼠,在3 组之间不存在显著差异;(5)在相互友好行为上,3 组间存在显著差异。其中社群2 中的雄性之间的友好行为频次最高,社群3 中次之,社群1 中最低。结果表明,雄性之间的攻击行为并不因为雌性的存在而增强,反而会可能减少,我们推测这可能因为雄性要花费更多的时间用于社群探究和交配,从而减少了雄性之间的斗争。     

青春期雌性根田鼠初次择偶行为与雄性优势等级

在实验室条件下，研究了性刚成熟的雌性根田鼠对一对互为同胎同胞的以体重划分为优势和从势的雄性个体的择偶行为，目的在于检验雄性社会等级是否影响其配偶选择。结果发现：(1)根田鼠存在显著的偏雄性二型，而雄性动物优势度与体重大小成正比；(2)在Y字型迷宫中，初次发情雌鼠对优势雄鼠的访问时间、社会探究频次和友好频次等显著大于从属雄鼠；(3)在随后24h的一雌二雄共居中，优势雄优先交配；从属雄的干扰不能中断优势雄鼠的交配。这些结果说明，体重差异可以作为划分雄性优势的指标，社会等级可以作为择偶行为的一个依据。优势雄鼠与处于从属地位的同胞相比，能获得优先交配机会。同胎同胞虽具有遗传相似性，但雄性竞争的结果表现在体重和社会等级上具有显著差异，表明配偶选择在发挥着作用。根田鼠是一雄多雌制，而且具有较大的偏雄性二型，提示社会等级、雌性选择和婚配制度有密切关联[动物学报49(3)：303—309，2003]。     

半野生麋鹿集群行为的初步研究

麋鹿在半野生状态下喜集群活动,不同月份集群的大小以及各种组群出现的频率均有所变化。半野生麋鹿有５种组群类型：雄鹿群、母仔群、混合群、雄仔群和仔鹿群。其中混合群出现频率最高,集群最大,是其主要的组群类型,雄鹿群最为稳定,母仔群中采食组群所占比例和组群大小都大于非采食活动的组群,雄仔群和仔鹿群出现频次较少。麋鹿发情期与非发情的集群有明显差异,不同生境内的集群也有所不同。     

四川短尾鼩尿液气味在社会等级识别中的作用

本文在中立竞技场中通过两两互作确定四川短尾鼩(Anourosorex squamipes)同性个体间的社会等级,并在此基础上利用其尿液,研究不同社会等级个体的自我/非自我识别能力及模式、尿液气味的行为响应机制,以及社会等级识别能力。结果表明：(1)四川短尾鼩优势个体表现攻击行为较多,从属个体防御行为较多,优势个体的标记行为显著高于从属个体。攻击行为表现为同等级雄性高于同等级雌性,且雌性间的攻击强度低于雄性。(2)从属个体和优势个体分别对自身尿液气味和非自身尿液气味存在明显偏好差异。不同性别、等级个体自我识别模式差异不明显,不同社会等级个体对于自身识别模式和非自身尿液的行为反应模式均不同。不同社会等级个体具有自我识别能力且能力不同。(3)四川短尾鼩能够识别不同社会等级个体的尿液气味,雌性对雄性尿液更感兴趣,雄性对优势雄性尿液选择回避。雄性对其他个体的访问时间与嗅舔频次均显著高于雌性,雌雄个体在识别不同社会等级的尿液气味时存在性二型。     

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.     

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.     

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.     

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.     

Seasonal changes of reproductive behavior and fecal steroid concentrations in Père David's deer.

From July 1997 to September 1997 and from March 1998 to July 1998, we studied reproductive behaviors of Père David's deer in Dafeng, China. During the field behavioral observations, we collected fresh voided fecal samples from the Père David's stags and hinds periodically and kept those samples under -20 degrees C until laboratory analysis. We analyzed the fecal testosterone, estradiol, and progesterone concentrations in those samples using radioimmunoassay. During this study, we also recorded 17 types of male reproductive behaviors and nine types of female reproductive behaviors. Reproductive behaviors and the fecal steroid concentrations showed overt seasonal fluctuations. There were statistically significant correlations between some male reproductive behaviors, such as anogenital sniffing, urine sniffing, urine spraying, wallowing, bellowing, antler adorning, Antler swags mud, chasing, herding hinds, chin resting, mounting and copulating, with the fecal testosterone concentrations. These results suggested that seasonal reproductive behaviors in stags are strongly associated with circulating testosterone. We also found that some female reproductive behaviors fluctuate corresponding with changes in fecal estrogen concentrations. Although there was no direct evidence to confirm the correlations between female reproductive behaviors and fecal estrogen in our experiment, we could not rule out that reproductive activities of hinds were largely related to ovarian estrogen secretion, and estrogen is necessary for inducing female reproductive behaviors.     

Seasonal Changes of Reproductive Behavior and Fecal Steroid Concentrations in Père David's Deer

From July 1997 to September 1997 and from March 1998 to July 1998, we studied reproductive behaviors of Père David's deer in Dafeng, China. During the field behavioral observations, we collected fresh voided fecal samples from the Père David's stags and hinds periodically and kept those samples under −20°C until laboratory analysis. We analyzed the fecal testosterone, estradiol, and progesterone concentrations in those samples using radioimmunoassay. During this study, we also recorded 17 types of male reproductive behaviors and nine types of female reproductive behaviors. Reproductive behaviors and the fecal steroid concentrations showed overt seasonal fluctuations. There were statistically significant correlations between some male reproductive behaviors, such as anogenital sniffing, urine sniffing, urine spraying, wallowing, bellowing, antler adorning, Antler swags mud, chasing, herding hinds, chin resting, mounting and copulating, with the fecal testosterone concentrations. These results suggested that seasonal reproductive behaviors in stags are strongly associated with circulating testosterone. We also found that some female reproductive behaviors fluctuate corresponding with changes in fecal estrogen concentrations. Although there was no direct evidence to confirm the correlations between female reproductive behaviors and fecal estrogen in our experiment, we could not rule out that reproductive activities of hinds were largely related to ovarian estrogen secretion, and estrogen is necessary for inducing female reproductive behaviors.     

A New Gene Mapping Resource: Interspecies Hybrids between Pere David's Deer (Elaphurus Davidianus) and Red Deer (Cervus Elaphus)

Three male F(1) hybrids between Pere David's deer and red deer were mated to red deer to produce 143 backcross calves. The pedigrees are a rare example of a fertile between evolutionarily divergent species. We examined the use of these families for genetic mapping of evolutionarily conserved (Type I) loci by testing for genetic linkage between five species-specific protein variants and 12 conserved DNA probes. Two probes were homologous, and the remainder syntenic, to the protein coding loci in cattle or humans. Using six restriction enzymes, each DNA probe detected one or more restriction fragments specific to Pere David's deer. Linkage analyses among the species-specific variants placed the loci into four linkage groups within which linkage between adjacent loci and gene order was supported by a LOD > 3. The linkage groups were (HPX, HBB)-FSHB- ACP2, LDHA-CD5-IGF2, BMP3- (GC, ALB)-(KIT, PDGFRA) and LDLR-C3-FGF1. Southern and protein analysis of LDHA and ALB provided identical segregation data. These linkage groups were consistent with the cattle gene map and provide new information for comparing the gene maps of ruminants, humans and mice. The deer hybrids are an important new resource that can contribute to the comparative analysis of the mammalian genome.     

麋鹿鹿角脱落、群主更替、产仔的年节律及其环境影响因子

本文在2014—2016年三个冬季(12月—翌年2月)收集了北京南海子麋鹿苑半散放麋鹿自然脱落的角,并观察和记录了2015—2017年发情期(5—9月)群主更替和2016—2018年产仔期(3—7月)麋鹿幼仔出生情况,结合2014—2018年年平均气温、季平均气温、月平均气温、年降雨量、雨季开始时间、种群密度等环境因子,对鹿角脱落、群主更替、产仔等繁殖特征的年节律及其环境影响因子进行了研究。结果表明：1)麋鹿鹿角脱落、群主更替、产仔的年节律均存在年际差异。2)鹿角脱落时间为12月开始,1月下旬或2月上旬结束。3)发情期为5月下旬或6月上旬开始,9月上旬结束;2015—2017年发情期时间有延长的趋势。4)产仔期为3月中旬或4月中旬开始,5月下旬或7月下旬结束。5)麋鹿鹿角脱落、群主更替、产仔的年节律存在明显的同步关系,其中鹿角脱落开始时间、鹿角脱落高峰期、鹿角脱落结束时间、第一次发情期开始时间、群主更替高峰期、产仔期开始时间、产仔高峰期、产仔期结束时间与前一年度比较均出现同步提前的现象。6)鹿角脱落年节律存在随着12月平均气温升高而提前的现象;产仔期开始时间和产仔高峰期存在随着前一年9月平均气温的升高而提前的现象。7)鹿角脱落年节律表现出随着年降雨量的增多而提前的现象;第一次发情期开始时间、群主更替高峰期的年节律表现出随着前一年度年降雨量的增多而提前的现象。8)麋鹿鹿角脱落、群主更替、产仔的年节律均不存在随着种群密度升高或降低而提前或延迟的现象。麋鹿繁殖特征的年节律是一个复杂的过程,受气候、营养、种群密度、纬度等环境因子的影响。     

Dominance, cortisol, and behavior in small groups of female cynomolgus monkeys (Macaca fascicularis)

The relationships among social rank, basal cortisol concentrations, and social behavior were assessed in adult female cynomolgus monkeys (Macaca fascicularis). Subjects were 157 unrelated, reproductively intact animals housed in 30 small groups. Rank determinations were made monthly. Blood samples were collected on two occasions, 4.5 and 7.5 months following initial group formation. Regular behavioral observations were conducted on a subset of animals over a period of 4 weeks, 9 months following group formation. Analyses revealed that serum cortisol values were significantly correlated across the two sampling periods, with no significant change in absolute values. While social rank was positively correlated across both samples, there was no relationship between rank and cortisol. However, dominant and subordinate animals did differ in the rates of performance of aggressive and submissive behaviors. These data suggest that social rank does not influence baseline serum cortisol in adult female cynomolgus monkeys, despite stability in measures of rank and cortisol and the presence of the usual behavioral differences between dominants and subordinates.