大山雀的婚外父权调查

婚配制度作为一种进化稳定对策是动物对某一环境包括种群内部环境适应的结果,在动物生殖过程中具有重要意义.大山雀(Parus major)是一种广域分布物种,研究显示广域分布物种的形态、生理、行为、生态特征及婚配制度在不同地区或种群间存在显著差异,因此,该文选择了分布于中国辽宁仙人洞国家级自然保护区的大山雀(P.m.minor)种群开展其婚配制度研究.野外共采集了22巢大山雀亲代和子代血液样本.结果如下:(1)从11个微卫星位点中筛选出了8个多态性较好的微卫星位点用于大山雀父权鉴定,在母本已知的情况下确定父权的准确率可达99.98％;(2)巢内父权鉴定结果显示,31.8％ (7/22)的巢包含婚外父权,8.12％ (16/197)的子代为婚外子代.与其他森林雀形目鸟类相比,大山雀婚外子代的比例明显偏低(＜10％),每个存在婚外父权巢的婚外子代比例各异(55.6％～9.1％),且无明显规律.     

杂色山雀的研究现状

杂色山雀Parus varius是一种分布区域狭窄的小型森林洞巢鸟类,为东亚特有鸟类,主要分布在中国、日本和朝鲜半岛,全球现有8个亚种.中国仅有2亚种,P.v.varius繁殖于辽宁东部山区,P.v.castaneoventris仅分布于台湾,均为当地留鸟;除P.v.castaneoventris以外,其他亚种在日本均有分布.本文根据国内外对该物种的研究情况,对其亚种分布、生态生物学、种群遗传结构及形态学等方面作了综述,以期为后续研究提供参考.     

杂色山雀的繁殖成功率

2004～2006年的3～7月,在辽宁省白石砬子国家级自然保护区,对杂色山雀(Parus varius)的繁殖及繁殖成功率进行了初步研究。结果显示,杂色山雀主要营巢于海拔400～900m的阔叶杂木林、针叶林及林缘地带;繁殖期在3～7月,其洞巢种类多样,筑巢期约15d;巢为碗状,巢结构的2/3由苔藓构成;窝卵数为6～8枚,平均(6·92±0·92)枚(n=13);雌鸟单独孵卵,孵化期为(14·00±0·00)d(n=10);育雏由雌雄鸟共同承担,育雏期为(17·50±0·58)d(n=4)。杂色山雀繁殖成功率为50·95%,繁殖力为2·22。人为干扰是造成卵和雏鸟损失的主要原因,占总损失的74·19%。     

杂色山雀亲代喂食与子代乞食间的行为

在育雏期,晚成鸟的子代一般都是由双亲共同来抚育,子代为了更好地存活,会用自己的方式竞争获得更多的食物和更好的生存空间,同时亲代也会根据子代的乞食信号来分配食物.2011年3～7月采用针孔摄像技术录制了杂色山雀(Parusvarius)育雏期巢内亲代与子代间的行为,统计了亲鸟站位、雏鸟站位、雏鸟乞食强度及亲鸟的喂食情况等数据.分析结果表明:(1)雌雄亲鸟在巢中的站位各有特点,雄鸟在整个育雏期都喜欢站在距离巢口较近的位置;雌鸟站位不太固定,前期离巢口相对较远,中期和后期离巢口相对较近;(2)雏鸟离亲鸟越近,乞食强度越大,获得食物的机会就越多;离亲鸟越远的雏鸟越不爱乞食,所以站位对雏鸟的食物获得影响最大;(3)雌鸟承担主要的育雏任务,喂食频率远大于雄鸟;(4)育雏期的不同阶段雏鸟乞食强度、亲鸟喂食频率变化很大:中期雏鸟乞食强度最大,亲鸟喂食频率最高,后期雏鸟乞食强度最弱;(5)整个育雏期雌性亲本没有表现出明显的偏爱行为,但雄性亲本在中、后期更偏爱体型大的雏鸟.可见杂色山雀子代的行为和体型大小影响着亲代的食物分配,亲代也会根据雏鸟日龄调整站位和喂食行为.     

杂色山雀指名亚种繁殖期的鸣声行为

2012年3～7月,对辽宁仙人洞自然保护区9巢18只杂色山雀(Parus varius varius)个体及其雏鸟的鸣声进行了录音,共获取了9种类型鸣叫(呼唤、警戒、报警、恫吓、驱逐、惊叫、喂食、雏鸟乞食、集群)和5种类型鸣唱.通过语图分析得出音节类型18种,频率范围为800 ～18 900 Hz.对杂色山雀不同个体鸣声特征参数的比较发现,鸣声的句子和音节时长在不同个体之间存在显著性差异,而最高频率、最低频率在不同个体间均无显著性差异.本研究实现了对杂色山雀繁殖期鸣声参数的量化,有助于进一步研究其繁殖行为.     

社会性单配制鸟类婚外父权的发生和影响因素

一雌一雄单配制鸟类中,雌性个体与配偶外雄性发生交配的行为称为婚外交配,继而导致了婚外受精产生婚外子代的现象称为产生了婚外父权。婚外父权广泛存在于鸟类中,针对其发生和影响因素已经成为了鸟类行为生态学研究的热点。本文收集了近十年社会性单配制鸟类婚外父权方面的研究文献,从婚外父权的发生及其影响因素两个方面综述了单配制鸟类婚外父权的研究进展。婚外父权发生原因的探讨主要包括：1、从两性的角度探讨雌雄两性在婚外行为中不同的进化繁殖策略。雄性策略旨在增加自身的繁殖输出;有关雌性策略则提出了确保受精假说、食物供给假说、遗传利益假说等,但目前尚存争议;2、在遗传利益假说中较常见的又分为3个假说：“优秀基因”假说、“遗传相容性”假说和“遗传多样性”假说,该三种假说是针对雌性从遗传方面获得的利益而提出的,不断有报道指出雌性配偶选择会被潜在的雄性遗传特性所影响;3、非遗传利益——母系效应影响婚外父权的进化。一些研究指出遗传质量参数,如体重、身体大小、存活率和免疫应答等方面可能会存在母系效应。婚外父权发生的影响因素这里主要指环境因素,包括繁殖同步性、繁殖密度、栖息地环境、产卵及孵化时机等。由于物种不同,受到环境压力不同,导致婚外父权发生率千差万别。最后本文针对未来的研究方向做出了展望。尽管近十年的研究进一步解释了鸟类婚外父权现象,但是该领域仍然存在并且产生了许多新的未解决的问题,而相关实验操作和理论的完善是深入探讨这些问题的关键。     

杂色山雀体型、子代性别比与鸣唱特征的相关性研究

鸣声在鸣鸟的生活史中具有重要作用,是其与外界进行信息交流最有效的方式之一。鸟类的鸣唱与其个体因素(如体型、子代性别比等)有着十分重要的联系。为了揭示个体体型、子代性别比与鸣唱特征之间的相关性,于2014—2015年3—6月在辽宁省仙人洞国家级自然保护区对杂色山雀指名亚种Parus varius varius的鸣声进行录制,测量了各体型参数,并对每巢雏鸟进行性别鉴定。结果显示:(1)杂色山雀体型与其鸣唱的最高频率具有显著相关性,但单一体型参数(体质量、跗蹠长度)与其鸣声参数相关性均不显著;(2)鸣唱语句长度与子代性别比具有显著相关性,子代性别比与其他鸣声参数均不存在显著相关性。本研究结果有助于进一步了解小型鸟类体型与鸣唱的关系,以及鸣唱与子代性别比之间的关系。     

繁殖期杂色山雀应激水平的皮质酮浓度 与繁殖投入的关系

皮质酮是鸟类重要的糖皮质激素,在其适应环境变化及压力应激反应中起重要的调节作用。非应激状态时,鸟类血浆皮质酮浓度处于基线水平,当鸟类面对应激刺激时,血浆皮质酮浓度迅速升高,应激水平的皮质酮对鸟类个体生存至关重要。然而,目前繁殖季鸟类血浆应激水平的皮质酮浓度变化及其与繁殖投入关系的研究结果存在种间差异,仍需在不同的物种中进行实验研究。本研究分析了繁殖季杂色山雀（Sittiparus varius）血浆应激水平的皮质酮浓度在繁殖阶段的变化,及育雏期亲鸟血浆应激水平的皮质酮浓度与繁殖参数和亲鸟育雏投入的关系。结果显示,与求偶期相比,育雏期杂色山雀亲鸟血浆应激水平的皮质酮浓度极显著升高（P < 0.001）,雌性与雄性亲鸟之间无显著性差异（P > 0.05）;雌性与雄性亲鸟血浆应激水平的皮质酮浓度与繁殖参数、亲鸟递食投入、巢防卫行为均无显著相关性（P > 0.05）。     

杂色山雀(Parus varius)亲代喂食与子代乞食间的行为交流

在育雏期,晚成鸟的子代一般都是由双亲共同来抚育,子代为了更好地存活,会用自己的方式竞争获得更多的食物和更好的生存空间,同时亲代也会通过子代的乞食信号来分配食物。2011年3月-7月采用针孔摄像技术录制了杂色山雀(Parus varius)育雏期巢内亲代与子代间的行为交流,统计了亲鸟站位、雏鸟站位、雏鸟乞食强度及亲鸟的喂食情况等数据。分析结果表明：(1)雌雄亲鸟在巢中的站位各有特点：雄鸟在整个育雏期都喜欢站在距离巢口较近的位置;雌鸟站位不太固定,前期离巢口相对较远,中期和后期离巢口相对较近;(2)雏鸟离亲鸟越近,乞食强度越大,获得食物的机会就越多;(3)杂色山雀主要是雌鸟担任育雏任务,喂食频率远大于雄鸟;(4)育雏期的不同阶段雏鸟乞食强度、亲鸟喂食频率变化很大：中期雏鸟乞食强度最大,亲鸟喂食频率最高,后期雏鸟乞食强度最弱;(5)整个育雏期雌性亲本没表现出明显的偏爱行为,但雄性亲本在中期表现出了偏爱大的雏鸟。     

杂色山雀对人工巢箱内苔藓类巢材的选择

2008年及2009年每年的3～8月,在辽宁省仙人洞国家级自然保护区对杂色山雀(Parus varius)人工巢箱内苔藓类巢材进行了收集和鉴定,发现巢中共有藓类植物7科9属。其中,小锦藓属(Brotherella)、青藓属(Brachytheccium)、棉藓属(Plagiothecium)及绢藓属(Entodon)所占比重较大。我们分别按照样方和苔藓群落采集了周围环境中的藓类植物,鉴定出7科8属的藓类植物;采集到35个苔藓群落,出现最多的是东亚小锦藓(B.fauriei)群落。结果显示,杂色山雀对苔藓的选择可能与苔藓的生物学特性、分布和数量有关。另外通过统计分析和比较得出,按照苔藓群落来采集的方式更适合本文的研究。     

Copulation behaviour and paternity in shy albatrosses (Thalassarche cauta)

We investigated the mating system of shy albatrosses Thalassarche cauta by combining behavioural observations during the pre-laying period with genetic paternity analysis. Genetic data on the mating systems of several procellariiform seabirds have recently become available, but data on the reproductive behaviour of these species are rarely obtained. Our main aims were to describe the copulatory behaviour of this species and identify how males achieve within-pair and extra-pair paternity (EPP). Most copulations occurred on the nest, were unforced and were within-pair. Females controlled the success of copulations and were observed soliciting extra-pair matings. Within-pair and extra-pair copulations were behaviourally similar. A low frequency (7–10%, n =29 chicks) of EPP was detected despite male use of frequent copulation as a paternity guard. The pre-laying foraging exodus of female shy albatrosses differed from that in other albatrosses: it was relatively short in length, lasting c . 2 days, and within-pair copulations occurred after the female's return 2 days before laying. This may reflect the close proximity of feeding grounds to the breeding colony.     

Reed bunting females increase fitness through extra-pair mating with genetically dissimilar males

Females of many socially monogamous species accept or even actively seek copulations outside the social pair bond. As females cannot increase the number of offspring with promiscuous behaviour, the question arises why they engage in extra-pair mating. We used microsatellite data to determine paternity, heterozygosity and genetic relatedness in the reed bunting (Emberiza schoeniclus), a species with high levels of extra-pair paternity (EPP). We found that extra-pair young (EPY) were more heterozygous than within-pair young (WPY). The high heterozygosity of the EPY resulted from a low genetic similarity between females and their extra-pair mates. EPY were heavier and larger when compared with their maternal half-siblings shortly before they left the nest. Recapture data indicated a higher fledgling survival of EPY compared with WPY. Our data suggest that reed bunting females increase the viability of their offspring and thus fitness through extra-pair mating with genetically dissimilar males.     

Subordinate male meerkats prospect for extra-group paternity: alternative reproductive tactics in a cooperative mammal

In cooperatively breeding species, subordinates typically suffer strong constraints on within-group reproduction. While numerous studies have highlighted the additional fitness benefits that subordinates might accrue through helping, few have considered the possibility that subordinates may also seek extra-group matings to improve their chances of actually breeding. Here, we show that subordinate males in cooperative meerkat, Suricata suricatta, societies conduct frequent extraterritorial forays, during periods of peak female fertility, which give rise to matings with females in other groups. Genetic analyses reveal that extra-group paternity (EGP) accrued while prospecting contributes substantially to the reproductive success of subordinates: yielding the majority of their offspring (approx. 70%); significantly reducing their age at first reproduction and allowing them to breed without dispersing. We estimate that prospecting subordinates sire 20-25% of all young in the population. While recent studies on cooperative birds indicate that dominant males accrue the majority of EGP, our findings reveal that EGP can also arise from alternative reproductive tactics employed exclusively by subordinates. It is important, therefore, that future attempts to estimate the fitness of subordinate males in animal societies quantify the distribution of extra-group as well as within-group paternity, because a substantial proportion of the reproductive success of subordinates may otherwise go undetected.     

Promiscuity, paternity and personality in the great tit

Understanding causes of variation in promiscuity within populations remain a major challenge. While most studies have focused on quantifying fitness costs and benefits of promiscuous behaviour, an alternative possibility--that variation in promiscuity within populations is maintained because of linkage with other traits-has received little attention. Here, we examine whether promiscuity in male and female great tits (Parus major)--quantified as extra-pair paternity (EPP) within and between nests--is associated with variation in a well-documented personality trait: exploration behaviour in a novel environment. Exploration behaviour has been shown to correlate with activity levels, risk-taking and boldness, and these are behaviours that may plausibly influence EPP. Exploration behaviour correlated positively with paternity gained outside the social pair among males in our population, but there was also a negative correlation with paternity in the social nest. Hence, while variation in male personality predicted the relative importance of paternity gain within and outside the pair bond, total paternity gained was unrelated to exploration behaviour. We found evidence that males paired with bold females were more likely to sire extra-pair young. Our data thus demonstrate a link between personality and promiscuity, with no net effects on reproductive success, suggesting personality-dependent mating tactics, in contrast with traditional adaptive explanations for promiscuity.     

Low historical rates of cuckoldry in a Western European human population traced by Y-chromosome and genealogical data

Recent evidence suggests that seeking out extra-pair paternity (EPP) can be a viable alternative reproductive strategy for both males and females in many pair-bonded species, including humans. Accurate data on EPP rates in humans, however, are scant and mostly restricted to extant populations. Here, we provide the first large-scale, unbiased genetic study of historical EPP rates in a Western European human population based on combining Y-chromosomal data to infer genetic patrilineages with genealogical and surname data, which reflect known historical presumed paternity. Using two independent methods, we estimate that over the last few centuries, EPP rates in Flanders (Belgium) were only around 1–2% per generation. This figure is substantially lower than the 8–30% per generation reported in some behavioural studies on historical EPP rates, but comparable with the rates reported by other genetic studies of contemporary Western European populations. These results suggest that human EPP rates have not changed substantially during the last 400 years in Flanders and imply that legal genealogies rarely differ from the biological ones. This result has significant implications for a diverse set of fields, including human population genetics, historical demography, forensic science and human sociobiology.