瓦屋山2种山雀的生态位分化和共存

黑冠山雀(Parus rubidiventris)和煤山雀(P.ater)在瓦屋山同域分布,生态位相似,是潜在的竞争者。在垂直分布上,煤山雀的生态位宽度为5.237,显著高于黑冠山雀的2.792,而在水平分布和活动基质2个维度上,二者的生态位宽度值相当。尽管3个维度上2种山雀生态位重叠值均大于0.7,但是它们对3个维度上的资源利用分别存在不同的偏好和侧重。2种山雀的身体量度除了喙宽外均有极显著差异,这应该是2种山雀对环境适应的结果。这2种山雀在巢址选择上各项参数差异均不显著,表明适宜的洞巢可能是它们种间竞争的主要对象。2种山雀的种群数量可能与居留类型有关。空间生态位或食物生态位的分离及适宜的种群数量和比例是它们得以共存的几个重要因素。     

杂色山雀的反盗食贮食策略

盗食现象在贮食动物中非常普遍,种内竞争者或种间竞争者的盗食是贮藏食物丢失的主要原因之一,同时也是贮食动物贮食行为不断进化的重要选择压力,开展反盗食贮食策略的研究有助于阐释贮食行为进化的原因和对贮食鸟类的保护。杂色山雀(Parus varius)秋季有明显的贮食行为,贮食方式为分散贮食。通过人为投食的方法,以投食点为中心,根据贮食点距投食点距离远近设定了被盗高风险区、被盗中风险区和被盗低风险区,研究杂色山雀个体在无潜在盗食者和有潜在盗食者的情况下,在上述3个风险区贮食比例的变化,探讨杂色山雀个体基于自身以及周围环境条件,在权衡取食及贮藏过程中时间及能量的花费和收益后,其反盗食贮食策略的选择。40只杂色山雀个体贮食策略的统计结果表明,不论潜在盗食者出现与否,杂色山雀个体贮食的总体趋势大致相同,即主要将食物贮藏在中风险区,而高风险区和低风险区贮食比例较低;比较种间和种内潜在盗食者出现的情况下与无潜在盗食者时,杂色山雀个体在各贮食风险区的贮食策略:种间及种内潜在盗食者的出现,都会引起被盗高风险区贮食比例降低(种间潜在盗食者:F=3.174,P0.05;种内潜在盗食者:F=90.475,P0.05),低风险区贮食比例上升(种间潜在盗食者:F=220.440,P0.05;种内潜在盗食者:F=15.651,P0.05);但种间潜在盗食者出现时,杂色山雀个体在被盗中风险区的贮食比例降低(F=143.749,P0.05),而种内潜在盗食者出现时,杂色山雀个体在被盗中风险区贮食比例不变(F=0.776,P0.05),即杂色山雀个体对种间潜在盗食者的出现更加敏感,防范盗食的投入更大。此外,分别比较无潜在盗食者、有种间潜在盗食者和有种内潜在盗食者存在的情况下,雌雄杂色山雀个体在各贮食风险区的反盗食贮食策略,发现其结果也明显不同。无潜在盗食者和种内潜在盗食者出现时,雄性杂色山雀为预防灾难性盗食发生而付出的努力较雌性个体高(无潜在盗食者:高风险区t=4.962,df=16.546,P0.05;中风险区t=﹣0.890,df=29.255,P0.05;低风险区t=﹣2.166,df=30,P0.05。有种内潜在盗食者:高风险区t=﹣0.152,df=29,P0.05;中风险区t=2.352,df=19.568,P0.05;低风险区t=﹣2.287,df=19.563,P0.05);种间潜在盗食者出现时,雌雄杂色山雀反盗食贮食策略趋于一致(高风险区t=1.361,df=29,P0.05;中风险区t=﹣0.194,df=21.529,P0.05;低风险区t=﹣1.599,df=29,P0.05)。     

食物和季节因素对杂色山雀贮食行为的影响

鸟类的贮食行为受很多因素的影响,其中食物和季节是两个十分关键的因子。采用人工投食的方式研究杂色山雀冬、春季节对松子和葵花籽两种食物的贮食选择,以及食物和季节因素对贮食位点的空间分布和微生境选择的影响。结果发现,杂色山雀优先贮食松子,仅在春季贮食少量葵花籽;主要选择树皮裂缝、灌木根部、草丛、石缝和苔藓下面五种生境进行贮食;其贮食位点的空间分布呈分散状态,集中于投食点100m范围内,密度分布随食物搬运距离的增加而递减。其贮食模式和贮食位点的微生境选择均受季节因素的影响,其中,贮食模式的季节变化可能是受生境中松子这种重要食物可获得性的下降所致,而杂色山雀冬季对树皮裂缝的利用率明显高于春季,可能是与冬季的积雪覆盖限制了其对地面贮食点的利用有关。杂色山雀的就近贮食模式可能是为了增加贮食效率,关于不同个体之间贮食位点的差异,以及季节变化对贮食位点的空间分布格局的影响还需进一步研究。     

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。     

杂色山雀繁殖期与非繁殖期食物组成及利用

食物资源是动物赖以生存的重要物质基础和能量来源。研究鸟类不同时期的食物选择和利用对于了解该物种取食行为的可塑性和生态适应性将具有重要意义。本文于2012年4月至2013年10月,通过取食行为观察、育雏分析和投食实验来研究杂色山雀(Parus varius)繁殖期与非繁殖的食物组成及利用的差别。结果发现,杂色山雀繁殖期与非繁殖期存在显著的食物差别,繁殖期完全取食动物性食物,而非繁殖期则以植物性食物为主,兼食少量昆虫。繁殖期成鸟取食的食物主要为鳞翅目、蜘蛛目、鞘翅目、双翅目以及少量的直翅目、异翅亚目以及膜翅目动物,育雏的食物资源主要为鳞翅目幼虫(67.86%),其次为蜘蛛目、鳞翅目成虫、膜翅目以及鞘翅目幼虫和少量直翅目,但与成鸟的食物存在显著区别,特别是鳞翅目幼虫的比例极高。非繁殖期的食物主要以植物种子(48.91%)和浆果为主(51.09%),其中,浆果的比例较高,对16种潜在食物资源的取食选择也证明对浆果类食物具有较明显的偏爱。杂色山雀不同时期的食物差别和食性的可塑性可能是由于食物供给和能量需求的季节性变化造成的,但对于自然生境中各种食物资源丰富度的季节性变化,及对杂色山雀取食选择的影响还需进一步研究。     

人工落叶松林中冬季鸟类混合群的相互关系研究

通过对人工落叶松林中冬季鸟类混合群的观察表明,主要由沼泽山雀、长尾山雀、普通鳾和旋木雀等四种鸟构成冬季混合群,并以沼泽山雀为主体。每种鸟在混合群中出现频次不同、取食生态位宽度和重叠不同。构成了对资源的分割,从而减少了竞争,使之能共存于混合群中。这种生态位的分化与形态特征有关,特别与嘴大小有关系。     

红头长尾山雀繁殖生态的研究

在安徽合肥紫蓬山区,对红头长尾山雀的繁殖生态进行了系统的观察,并研究了雏鸟的生长特征。研究表明,红头长尾山雀自3月中旬以后开始发情,3月下旬开始营巢,巢位于杉树枝下,球形侧开口或侧顶开口。4月初产卵,每窝平均产6．25枚卵,孵化期为15—16d,孵化率为88％。育维期15—16d,22维全部成活到离巢,雏成活率达100％。体重、翅、嘴峰、跗跖、第5枚初级飞羽和尾羽等形态学参数适合用Logistic曲线方程拟合,拟合参数中,嘴峰的拐点最小,飞羽、翅和尾羽的增长率较高,表明与取食有关的形态优先发育,与飞行有关的形态则在后期发育较快。     

大山雀和褐头山雀种间关系研究

在地理分布重叠地区的大山雀Parus major和褐头山雀Parus montanus栖息地海拔高度不同。大山雀栖于海拔2,370米以下,集中于次生杨桦林和老年人工杨树林;褐头山雀栖于海拔2,300米以上,集中于混交林和山杨栎林。它们的栖息地有一定重叠,但重叠值较低。重叠地带觅食生态位的四维中,树种选择一维有所不同,其余三维生态位重叠值均较大。两个种营养生态位有一定重叠,食物大小的重叠值更大。在进化发展过程中,两种山雀间完全的生态分离没有发生。     

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.     

人工巢箱条件下两种山雀鸟类的同域共存机制

种间竞争会导致鸟类对自身的生存策略进行多方面调整,将多种因素结合起来分析不同鸟类的同域共存机制具有十分重要的意义.2009-2011年的3-7月,在辽宁省东部山区以悬挂人工巢箱的方法招引杂色山雀(Parus varius)和大山雀(Parus major),通过比较这两种鸟类在繁殖时间、繁殖参数和巢址选择上的差异,分析其繁殖期的栖息地选择及繁殖对策,探讨这两种鸟类同域共存的机制.结果表明,生态位的部分分离是大山雀和杂色山雀能够长期共存的基础.两者都是在繁殖期到来立即开始繁殖,均出现两次繁殖高峰,采用消减窝卵数的对策来适应环境质量的下降.对繁殖参数的分析表明,两种鸟类的繁殖对策都为k-选择,但大山雀略偏向r端,大山雀倾向高窝卵数、低繁殖成功率的繁殖策略;杂色山雀倾向低窝卵数、高繁殖成功率的繁殖策略.两种鸟类都选择在林龄较长的森林中筑巢,但是大山雀较注重巢址的安全性能,巢距地面较高,周围乔木密集;杂色山雀较为注重植被类型的选择,巢址多选择在植被多样、灌木生长茂盛的针阔混交林中,推测可能与杂色山雀的食物构成有关.     

山地次生林冬季鸟类集群的初步研究

通过山地次生林两冬季鸟类集群的研究表明,集群的鸟类主要是由大山雀、沼泽山雀和普通Ｓｈｉ组成;每群个体数为７．４１－１０．６３只;据群的行为（单种群、多种集群和单独个体）分析,大多数种类是单独个体活动,冬季鸟类集群大小和组成有空间的变化。气候因子影响群的形成和行为。     

DIFFERENTIATION OF FORAGING NICHE AMONG TITS, PARUS SPP., IN NORWAY DURING WINTER

The winter foraging behaviour of Willow Tit Parus montanuss, Coal Tit P. ater and Crested Tit P. cristatus was studied in several spruce-dominated, coniferous forest areas in which either Willow Tit alone, Willow Tit and Coal Tit, or all three species together were present. In the areas where Crested Tits were absent, Willow Tits foraged significantly more often in the outermost parts of spruce branches and less frequently on the parts of branches closer to the trunk, than in the area in which Crested Tits were present. In the area in which all three species were present, the same change in foraging pattern of Willow Tits was found in mixed flocks in which Crested Tits were lacking. The foraging pattern of Coal Tits was unaffected by the presence of members of either or both the other two tit species. It is suggested that the expansion of the feeding niche of Willow Tit in the absence of Crested Tit is a result of ecological release, due to a reduction in the degree of interspecific competition.     

Factors influencing Willow Tit Poecile montanus site occupancy: a comparison of abandoned and occupied woods

The British Willow Tit Poecile montanus kleinschmidti underwent a decline of 85% between 1970 and 2003. The cause of this decline is unknown. However, several hypotheses have been put forward to account for it: competition from other tit species, predation by Great Spotted Woodpeckers Dendrocopos major and habitat change. In order to test these, woods that are currently occupied by Willow Tits were paired with woods (within 50 km) that had been abandoned by Willow Tits five or more years previously. Point counts for other tit species (potential competitors) and woodpecker species (potential predators) were carried out at ten evenly spaced points throughout each wood. Habitat variables were collected within a 50-m radius of where a Willow Tit was located (in the occupied woods) or where maps showed a Willow Tit had been located (for abandoned woods). No evidence was found for differences in numbers of potential competitor or potential predator species in abandoned and occupied woods. Soil water content was found to be higher at occupied sites. No other habitat features differed between the two categories of site. The drying up of British woods could therefore be implicated in the Willow Tit decline and this warrants further investigation.     

STRUCTURE AND FORAGING PATTERNS OF FLOCKS OF TITS AND ASSOCIATED SPECIES IN AN ENGLISH WOODLAND DURING THE WINTER

Blue Tits were the commonest and most frequent members of mixed-species insectivorous flocks during the winter at Wytham Wood, Oxford. Six common flocking species (Blue Tit, Longtailed Tit, Great Tit, Coal Tit, Marsh Tit and Golderest) made up nearly two-thirds of the birds in the study area. A tentative interspecific social dominance hierarehy was constructed: Great Tit > Blue Tit > Marsh Tit > Coal Tit > Long-tailed Tit and Goldcrest. Blue Tits permitted conspecifics to approach them more closely than did other flock members. Blue Tits most frequently led the flocks, but no more than would be predicted by their abundance. Great Tits, and Marsh Tits and Coal Tits to a lesser extent, defended territories during the winter. Two of the six common species frequented the inner parts of branches (Great Tit, Coal Tit), two the outer parts of branches (Blue Tit, Marsh Tit), and two the twigs (Long-tailed Tit, Goldcrest). Members of each of these species-pairs showed marked differences in height of foraging and/or species of tree frequented. Because of their abundance, the impact of Blue Tits outside of their most highly frequented foraging zones may exceed that of species concentrating in these other zones (e. g., twigs high in trees). The Coal Tit foraged most diversely, the Great Tit least diversely. Species that foraged diversely in one of the three foraging categories (species of tree, substrate, height) usually foraged less diversely than most other species in the other dimensions. Species usually overlapped least in the zones exploited (inner parts of branches, etc.). Great Tits overlapped with other species less than did any other common flock member, and Coal Tits were the next lowest in this regard.     

Effects of habitat fragmentation on foraging behaviour of tits and related species: does niche space vary in relation to size and degree of isolation of forest fragments?

We studied the winter foraging niches of tits and related species in deciduous forest fragments varying in size between 1 and 30 ha (plus one forest of 200 ha) in order to investigate the influence of forest fragmentation on foraging niches Very few correlations between niche structure (foraging niche, width and overlap) and forest size or isolation turned out to be significant This implies that either the species that disappear in small fragments are those that suffer most from competition (making the effect unmeasurable), or that competition is relatively unimportant for niche structure In any case we find no evidence that foraging niches are strongly affected by the changes (in habitat and/or community structure) associated with fragmentation     

THE SELECTION OF TITS PARUS SPP. BY SPARROWHAWKS ACCIPITER NISUS

Selection of the Wytham Wood population of tits by Sparrowhawks during the tit post-fledging period was studied over three years. Within three weeks of the median date of tit fledging, juvenile Blue Tits were selected more frequently than juvenile Great Tits but thereafter juvenile Great Tits were selected more frequently; it is suggested that the former was due to greater vulnerability of Blue Tits in the two to three weeks after fledging, and the latter due to changes in availability due to immigration/emigration of juvenile tits. When juvenile tits killed by hawks were compared on the basis of brood and physical characteristics with their cohorts in the entire population and in those surviving to the following year, only one factor, fledging date, was found to have affected selection by hawks. It is believed that this was because hawks selected late fledged young still in family parties in preference to early fledged young which had already become independent of their parents and were foraging in the better cover of the forest canopy. When adults killed by hawks were compared with all adults available for selection on the basis of species, sex, timing of nesting and whether or not they had been born in the wood, the only selection trend found was that male Great Tits were taken more often than females due, possibly, to greater risk of exposure for males while foraging or during territorial behaviour. When compared with availability, juveniles were selected more frequently than adults in only one year, possibly a result of yearly differences in the number of prey available per hawk, the rate of non-predator related juvenile mortality or unknown hawk hunting strategies. The overall finding of relative non-selectivity by hawks was attributed to the surprise factor associated with the hunting methods used by Sparrowhawks in woodland.     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

Food resource matching by foraging tits Parus spp. during spring-summer in a Mediterranean mixed forest; evidence for an ideal free distribution

We analysed whether patterns of microhabitat use by Blue Tits Parus caeruleus , Great Tits Parus major and Crested Tits Parus cristatus inhabiting a mixed forest consistently matched the patterns of food availability experienced by foraging birds during spring-summer. The use of five microhabitats by each bird species (the foliage of three tree species, shrubs and ground) and the availability of food in trees during the prebreeding, breeding and post-breeding periods of the birds' annual cycle were measured. All three tit species foraged mainly in the outer part of tree canopies (small branches and leaves or needles). Tit distributions between tree species matched food resource distributions irrespective of overall food resource levels, which varied four-fold between the study periods, and tit species. Tits also exploited secondary microhabitats (shrubs and ground) in periods of low food availability; Blue Tits tended to use shrubs, whereas Great and Crested Tits foraged on the ground. Between-trees distributions fitted that expected from an ideal free distribution, suggesting that food availability and intraspecific exploitative competition were the main factors governing tree use by tits. In contrast, patterns of use of secondary microhabitats (shrubs and ground) seemed to indicate a role for the species-specific morphological configurations of each tit species since Blue Tits are better adapted to hang and tended to forage in shubs, whereas Great and Crested Tits are better adapted to feed on horizontal surfaces and tended to forage on the ground. No evidence of interspecific interactions was observed. Overall, the results pointed to an independent exploitation of Mediterranean mixed forest by each bird species, food availability and food accessibility being the main factors affecting microhabitat use by foraging tits.     

Seeing without being seen: a removal experiment with mixed flocks of Willow and Crested Tits Parus montanus and cristatus

This paper tests the hypothesis that foraging site selection reflects a trade-off between the various needs for concealment from predators, to find food, and for the individual to maintain some view of its surroundings. After removal of Crested Tits Parus cristatus (the dominant species in mixed flocks), Willow Tits P. montanus did not decrease their foraging heights as expected but remained in the most exposed parts of young pines. In contrast, after removal of Willow Tits, Crested Tits increased their foraging height from the sheltered lower canopy to sites previously occupied by Willow Tits. When flock size was reduced, the birds maintained the same high levels of vigilance without concealing themselves in dense vegetation. I suggest that flock members may benefit from foraging in sites that afford good anti-predator vigilance.