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。     

杂色山雀双亲差异性育雏策略

社会性单配制鸟类的配偶双方在抚育子代时常存在性别差异,不同鸟种的雌雄双亲往往采取不同的育雏策略。以杂色山雀(Sittiparus varius)为研究对象,2017年3—7月对繁殖巢箱进行录像监测,记录杂色山雀育雏期亲代投入情况。分析结果显示:1)双亲递食率在育雏前期(4—6日龄)无显著差异,而育雏后期(10—12日龄)雌性的递食率显著高于雄性。2)雌性亲鸟后期递食率较育雏前期显著增加;而雄性亲鸟育雏前期和后期递食率无显著差异。3)雌性递食率与自身喙宽呈极显著正相关,雄性递食率与双亲体征参数均无相关关系。总的来说,在育雏阶段,杂色山雀雌性亲鸟的递食率随着雏鸟的需求和自身身体质量发生调整,雌性在育雏后期递食率显著升高,而雄性亲鸟递食率无变化,这可能与育雏期双亲投入分工不同有关。     

乌鸫的繁殖行为与坐巢行为初步观察

于2009年3～7月、2010年3～6月,采用焦点动物观察法和全事件行为记录法对乌鸫(Turdusmerula)的孵卵及育雏行为进行了研究。结果表明,乌鸫是雌鸟孵卵,在孵卵期出现卵损失现象,具补卵行为;孵卵前期与后期的坐巢行为存在差异,且其坐巢时间、频次和坐巢率有随孵卵数递增的趋势。双亲育雏但以雌鸟为主,喂食模式3种:雌鸟喂食、雄鸟喂食、雄鸟将食物递给雌鸟由雌鸟喂食。随着雏鸟日龄的增长,喂食次数增多,暖雏行为减少,至育雏后期未观察到暖雏行为。为权衡孵卵期和育雏前期的能量分配,乌鸫在孵卵期及育雏前期分别采取时间长而频次少和时间短而频次多的坐巢策略。     

一对人工巢箱内繁殖的白眉姬鹟育雏习性研究

2011年6月10～24日,在北京松山自然保护区采用全日观察法,对在游览道路旁的人工巢箱内繁殖的1对白眉姬鹟的育雏行为及其栖息地环境条件进行了系统观察与数据采集.结果显示育雏前期和后期雌雄鸟日喂食次数及各时段雌雄鸟进出巢的平均次数均有一定差异,白眉姬鹟在育雏过程中雌鸟喂食次数随雏鸟生长而增加,雄鸟则无明显变化.游客及人为干扰对雌雄鸟出入巢穴均有一定影响,雌鸟较之雄鸟更为胆怯和谨慎.表明白眉姬鹟雌雄鸟在育雏中的行为存在分工和差异.     

秃鹫巢内育雏期亲鸟与幼鸟行为及时间分配

2016和2017年,采用焦点动物取样法及全事件记录法,在新疆和静县巴仑台镇研究了4个繁殖巢的秃鹫(Aegypius monachus)繁殖行为。通过红外相机和人工观察,构建了秃鹫巢内育雏期的行为谱,将秃鹫亲鸟的行为划分为9类33种,将雏鸟行为划分为6类28种。结果表明,亲鸟喂食次数的最高峰出现在12:30~13:30时,随后在15:30~18:30时之间出现一个小高峰。在育雏期,亲鸟行为以护幼、观望和警戒为主,雏鸟则以休息和观望为主。将巢内育雏期分为三个阶段:育雏前期(4~5月)、育雏中期(6月)、育雏后期(7月),运用单因素方差分析(one-way ANOVA)检验不同育雏阶段亲鸟、雏鸟行为时间分配的差异。结果显示,育雏前期与后期之间亲鸟的行为时间分配差异不显著(P0.05),前期与中期和中期与后期之间,亲鸟的行为时间分配差异均显著(P0.01);育雏前期、中期与后期的雏鸟行为时间分配差异均显著(P0.01)。国内秃鹫繁殖主要面临食物短缺、人类活动干扰等威胁。     

陕西红碱淖遗鸥育雏行为和雏鸟生长

2012年5～7月,应用e-Science信息技术和标记法,对陕西神木县红碱淖(N 38°13′～ 39°27′,E 109°42′～110°54′)遗鸥(Larus relictus)的育雏行为和雏鸟生长发育进行了研究.结果表明,雏鸟由双亲共同承担喂食.育雏前期,亲鸟采取直接喂食、食物呕吐于巢边和在巢中间断性喂食这3种喂食模式;亲鸟昼间平均喂食(0.706±0.036)次/h,夜间平均喂食(0.469±0.024)次/h,双亲在喂食频次上无显著差异(F=32.54,P＞0.05).育雏后期,主要采取双亲直接喂食和亲鸟把食物呕吐于地面上,由雏鸟自己取食的喂食模式;亲鸟昼间平均喂食(0.416±0.021)次/h,夜间平均喂食(0.331±0.018)次/h,亲鸟喂食次数与雏鸟的日龄存在相关性(r =0.074,P＜0.05).随着雏鸟日龄的增长,暖雏次数趋于减少,而在炎热晴天、降雨和大风等天气状况下,暖雏时间和护雏行为都增强.雏鸟20日龄后未再观察到暖雏行为.雏鸟体长及外部器官的形态学参数适合用Gompertz曲线方程拟合.同时,与其近缘种黑嘴鸥(L.saundersi)的育雏行为和雏鸟生长进行了比较.     

不同性别杂色山雀血浆活性氧和超 氧化物歧化酶与繁殖成效的关系

氧化应激是指活性氧等氧化剂的产生大于抗氧化防御系统清除能力时的不平衡状态,是个体应对内外环境刺激的适应性生理机制,是衡量个体身体状况的综合性生理指标。为探究氧化应激对鸟类繁殖的预测作用,本研究测量了育雏前期(雏鸟6～8日龄)杂色山雀(Sittiparus varius)亲鸟血浆氧化应激分子活性氧和超氧化物歧化酶含量,通过巢箱监测获得了杂色山雀的繁殖参数,采用偏最小二乘回归法分析杂色山雀氧化应激对其雏鸟出飞率和繁殖成功率的预测作用。结果显示,雌性杂色山雀亲鸟血浆活性氧浓度与雏鸟出飞率(n=13,P 0.05)、繁殖成功率(n=13,P 0.01)均呈显著负相关关系,即血浆活性氧水平越低的雌性杂色山雀,其繁殖巢雏鸟出飞率和繁殖成功率越高;雌性杂色山雀亲鸟血浆超氧化物歧化酶浓度与雏鸟出飞率(n=13,P0.05)、繁殖成功率(n=13,P 0.05)无显著相关关系。本研究中这两项血浆氧化应激标记物与杂色山雀雄性亲鸟繁殖成效间均无显著的相关关系。该结果表明,杂色山雀雌性亲鸟活性氧水平对其繁殖成效具有显著的预测作用。     

怒江河谷栗喉蜂虎的繁殖行为观察

2007年3月26日—7月16日,采用典型野外记录法和所有事件取样法对云南省保山市道街怒江河谷栗喉蜂虎繁殖行为进行了观察。结果表明：1）产卵期雄性栗喉蜂虎求偶喂食的食物以蜻蜓目和膜翅目昆虫为主,占83.56%;雌鸟接受喂食的可能性为96.83%,接受喂食后仅有17.49%的雌鸟与雄鸟进行交配。2）在孵卵期昼间,雄鸟每次坐巢的时间为（23.521.6）min,每日坐巢的时间占54.4%;雌鸟每次坐巢的时间为（25.811.5）min,每日坐巢的时间占42.1%,雌雄鸟每次坐巢的时间无明显差异。3）幼雏出壳后亲鸟对幼雏的暖雏时间逐日减少,至第20 d亲鸟不再暖雏。栗喉蜂虎的育雏期为29～33 d,在繁殖过程中帮助喂食的鸟确实能提高被助鸟的繁殖成功率。     

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.     

杂色山雀的繁殖生态

2004—2006年3—7月在辽宁省白石砬子国家级自然保护区对杂色山雀的繁殖生态进行了观察研究。结果表明:该鸟繁殖期为3—7月,筑巢地除了树洞之外,还见在墙缝、石缝、金腰燕旧巢及电柱孔洞中筑巢,筑巢主要由雌鸟完成,雄鸟从旁协助;产卵期为5～8d,日产1枚,窝卵数为(6.92±0.92)枚(n=13);产满窝卵数后即开始孵化;孵化由雌鸟单独完成,孵化期(14.00±0.00)d(n=10),坐巢时间(570.00±11.02)min.d-1(n=9)(不含夜间),平均坐巢时间(57.00±6.75)min.次-1(n=9),雄鸟担任警卫任务,孵化期较为敏感,受到干扰易弃巢;育雏期(17.50±0.58)d(n=4),由雌雄鸟共同喂雏,雏鸟出飞后先是在亲鸟的带领下在巢周活动,之后活动范围逐渐扩展,2～3d后基本离开巢区。     

Parental allocation of food to nestling tree swallows: the influence of nestling behaviour, sex and paternity

Parents are expected to invest more in young that provide the greatest fitness returns. The cues that parents use to allocate resources between their offspring have received much recent attention. In birds, parents may use begging intensity, position in the nest or nestling size as cues to provision the most competitive young or those most likely to survive. It may also benefit parents to invest in young differentially by sex or relatedness if the fitness returns of sons and daughters differ or broods are sired by multiple males. We examined the allocation of food to tree swallow, Tachycineta bicolor, nestlings in relation to their begging behaviour, size, sex and paternity. Provisioning by parents was not related to nestling size, sex or paternity. The begging behaviour of nestlings did not differ with respect to sex or paternity. Both parents were more likely to feed nestlings that begged first or were closer to the nest entrance, suggesting that parents allocate food resources in response to cues that nestlings control. As a consequence, brood reduction was facilitated by biased provisioning within the brood in addition to the nestling size hierarchies created by hatching asynchrony. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Features of begging calls reveal general condition and need of food of barn swallow (Hirundo rustica) nestlings

Altricial offspring of birds solicit food provisioning by complexbegging displays, implying acoustic and visual signals. Differentcomponents of begging behavior may function as reliable signalsof offspring state and thus reproductive value, on which parentsbase optimal parental decisions about allocation of criticalresources (e.g., food). We experimentally manipulated componentsof general condition of nestling barn swallows (Hirundo rustica)by (1) altering brood size by cross-fostering an unbalanced number of nestlings between pairs of synchronous broods andthus manipulating the level of within-brood competition forfood, (2) injecting some nestlings with a harmless immunogen,simulating an infection, and (3) preventing part of the nestlingsfrom receiving food for a short period while establishing controlgroups. We recorded rate of begging response by individual nestlings as parents visited the nest and recorded begging calls usinga DAT recorder to analyze six sonagraphic features of vocalizations.Our factorial experiment revealed that nestlings deprived offood begged more frequently when parents visited the nest comparedto their non—food-deprived nest mates. Food deprivationincreased duration of syllables forming begging calls, whereas brood size enlargement resulted in increased latency of responseto parental calls. Heavy nestlings in good body condition vocalizedat a relatively low peak frequency. To our knowledge, thisis the first study in which begging rate and sonagraphic structureof begging calls are shown to reliably reveal a diverse setof components of offspring general state, on which parental decisions may be based.     

Nest mate size, but not short-term need, influences begging behavior of a generalist brood parasite

Hosts of generalist brood parasites often vary with regardsto their life-history traits, and these differences have thepotential to influence the competitive environment experiencedby brood-parasitic nestlings. Although begging by brood parasitesis more exaggerated than their hosts, it is unclear if generalistbrood parasites modulate their begging behavior relative tohost size. I examined the begging behavior of brown-headed cowbird(Molothrus ater) nestlings when competing against nest matesthat differ in size and under different levels of short-termneed. Cowbird nestlings begged on nearly all feeding visits,responded to adults as fast as (or faster than) their nest mates,and typically begged more intensively than their nest mates.Latency to beg, time spent begging, and maximum begging postureof cowbirds were similar during supplementation and deprivationtreatments, indicating begging intensity was not influencedby short-term need. Time spent begging by cowbirds varied amonghosts of 3 different sizes when short-term need was standardized,suggesting that nest mate size strongly influenced begging behavior.Cowbirds obtained more food when competing against an intermediate-sizedhost due to lower provisioning rates of small hosts or becauseof increased competitive ability of large host nestlings. Overall,cowbirds obtained the greatest volume of food per unit timespent begging when competing against intermediate hosts, butthis value approached that of the small host when adjusted formodal brood size. These results demonstrate that cowbirds adjusttheir begging relative to the size of the hosts against whichthey compete but not to levels of short-term need.     

Corticosterone Promotes Scramble Competition Over Sibling Negotiation in Barn Owl Nestlings (Tyto alba)

In species with parental care, siblings compete for access to food resources. Typically, they vocally signal their level of need to each other and to parents, and jostle for the position in the nest where parents deliver food. Although food shortage and social interactions are stressful, little is known about the effect of stress on the way siblings resolve the conflict over how food is shared among them. Because glucocorticoid hormones mediate physiological and behavioral responses to stressors, we tested whether corticosterone, the main glucocorticoid in birds, modulates physical and vocal signaling used by barn owl siblings (Tyto alba) to compete for food. Although corticosterone-implanted (cort-) nestlings and placebo-nestlings were similarly successful to monopolize food, they employed different behavioral strategies. Compared to placebo-nestlings, cort-individuals reduced the rate of vocally communicating with their siblings (but not with their parents) but were positioned closer to the nest-box entrance where parents predictably deliver food. Therefore, corticosterone induced nestlings to increase their effort in physical competition for the best nest position at the expense of investment in sib?Csib communication without modifying vocal begging signals directed to parents. This suggests that in the barn owl stress alters nestlings?? behavior and corticosterone could mediate the trade-off between scramble competition and vocal sib?Csib communication. We conclude that stressful environments may prevent the evolution of sib?Csib communication as a way to resolve family conflicts peacefully.     

An immunological cost of begging in house sparrow nestlings

Parent–offspring conflict predicts that offspring should demand a greater parental investment than is optimal for their parents to deliver. This would escalate the level of offspring demand ad infinitum, but most of the models on the evolution of parent–offspring communication predict that begging must be costly, such costs limiting the escalation and defining an optimal level of begging. However, empirical evidence on this issue is mixed. A potential begging cost that remains to be accurately explored is a decrease in immunocompetence for offspring begging fiercely. This study experimentally analyses this cost in house sparrow (Passer domesticus) nestlings. A group of nestlings was forced to beg fiercely for a prolonged time while a control group begged at low levels, both groups receiving the same quantity of food. At the same time, the nestling response to an antigen (phytohaemagglutinin) was measured. Nestlings forced to beg fiercely showed a reduction in immunocompetence with respect to control chicks, but the two groups showed no difference in growth rate. The largest and the smallest nestlings in each brood showed a similar response to the treatment. These results strongly suggest a trade-off between begging and immunocompetence in this species. This trade-off may be a consequence either of resources from the immune system being reallocated to begging behaviour, or of adaptive immunosuppression in order to avoid oxidative stress. Steroid hormones are proposed as mediators of such a trade-off.     

Great spotted cuckoos respond earlier to the arrival of feeding foster parents and perform less erroneous begging when hungry than their magpie host nest‐mates

In many bird species, parents usually feed the first nestling that starts to beg before its nest‐mates. The pressure to avoid missed feeds could trigger nestlings to perform in erroneous begging in absence of parents, which has the same costs as begging in the presence of parents but without any reward. So, nestlings should try to minimize both erroneous begging and missed feeds simultaneously. The threshold to start begging is predicted to be lower for hungry nestlings and for nestlings that are unrelated to their nest‐mates, because they suffer lower inclusive fitness costs when depriving nest‐mates of food. In line with this idea, we found that brood parasitic great spotted cuckoo nestlings responded sooner than their magpie nest‐mates when an adult arrived to the nest. Under laboratory conditions, nestlings of both species rarely incurred in erroneous begging when food was abundant, but under conditions of restricted food, magpie nestlings increased erroneous begging while cuckoo nestlings did not. Highly conspicuous begging in cuckoos results in an increased predation risk, which could have resulted in stronger selection pressures on cuckoos to avoid erroneous begging, probably resulting in better developed perceptual abilities, allowing cuckoos to perform better than their host nest‐mates.     

Why brown-headed cowbirds do not influence red-winged blackbird parent behaviour

Brown-headed cowbirds, Molothrus ater, frequently parasitize red-winged blackbirds,Agelaius phoeniceus . The presence of a brood parasite, unrelated to both host nestlings and parents, has provoked speculation regarding within-brood food allocation and parental provisioning. This study is the first to compare directly the effect of brood parasitism on host parent and offspring behaviour in younger and older broods. We videotaped 28 unparasitized red-winged blackbird broods and compared them to 22 parasitized broods. Red-winged blackbird nestling begging appears largely unaffected by cowbird parasitism. The presence of the cowbird in the nest affected neither the latency nor duration of host nestling begging, but stimulated more frequent begging by red-winged blackbird nestlings following food distribution. Begging by cowbirds was unique in two ways: (1) cowbirds maintained a consistent begging effort throughout the nestling period (but did not receive a consistent food share); and (2) cowbirds begged longer and more frequently following the allocation of food. Persistent begging by the cowbird following the allocation of food has implications for the division of parental care, if by doing so the brood parasite is able to provoke the foster parent to increase provisioning, at the expense of brooding. We found no evidence for the adjustment of parental care. Neither the foraging rates nor the lengths of the parental feeding visits differed markedly between parasitized and unparasitized broods. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Nestling tree swallows (Tachycineta bicolor) alter begging behaviour in response to odour of familiar adults,but not their nests

Communication using chemical cues is important for many taxa, including birds, but the use of olfaction for intraspecific communication has been investigated only recently in passerines and is understudied in nestlings. To address this knowledge gap, we explored whether nestling tree swallows (Tachycineta bicolor) would recognize and respond to chemical cues of conspecifics, specifically testing begging responses to familiar and unfamiliar nest and adult odours. For the nest odour experiment, nests were treated with either orange essential oil or distilled water to create scented and unscented (control) odour environments, respectively. For the adult odour experiment, adults attending the nestlings were considered “familiar adults” and adults attending a different brood in the population were considered “unfamiliar adults.” We found that begging responses of nestlings did not differ in response to orange oil odour or water, but nestlings begged significantly longer and more intensely in response to odours of a familiar than an unfamiliar adult, regardless of adult sex. This provides evidence that tree swallows use chemical cues to alter their behaviour and opens up many exciting avenues of future research.     

Signalling of Nutritional Need by Magpie Nestlings

The relationship between begging behaviour, chick nutritional state, and parental distribution of food within broods was studied in 4- and 5-chick magpie Pica pica broods under natural conditions. Three components of the begging display (duration, latency, and posture) were highly correlated with each other and also with the emission and duration of begging calls. Begging performance was strongly influenced by the food intake of nestlings during the preceding 1-h interval, indicating that begging may reliably reflect the nutritional need of nestlings. Daily growth during the preceding day, as well as average cumulative food intake by the brood during the preceding 24 h, seemed not to affect begging in a similar way. Begging signals employed by hungrier nestlings involved a higher degree of muscular activity, thus supporting the prediction that nestlings in greater need should employ more costly signals. Overall, those nestlings who begged more tended to obtain more food, but the relationship between feeding success and begging behaviour was weak due to a high variation between broods in the way that parents seemed to respond to variations in begging behaviour. Possible causes for this variation, and its implications for the evolution of reliable begging displays, are discussed.     

Begging in the absence of parents by nestling tree swallows

Begging by nestling passerine birds has become a model systemfor studies in animal communication. Although most beggingoccurs when parents arrive at the nest to feed (here called"primary begging"), it also occurs between feeding visits andimmediately after parents leave the nest. Begging in thesecontexts (here called "secondary begging") may have relativelylittle influence on the probability of receiving food, but could increase the overall cost of the signal and thus influence nestlingbegging strategies. The purpose of our study was to determinehow often tree swallow (Tachycineta bicolor) nestlings begin contexts other than to parents with food and to examinewhat factors influence the frequency of this begging. Secondarybegging ranged from 7% of measured begging responses at day2 to 30% by day 8 and was more frequent when the interval betweenparental feeding visits was relatively long and when the timeto respond to the arrival of parents with food was short. Increasesin both age and intervisit interval were associated with decreasesin nestling response times, suggesting that secondary beggingmay be related to the speed with which nestlings respond to stimuli. We discuss possible functions of secondary beggingand raise the possibility that it may, in fact, be an error.