弄岗熊猴的活动节律和活动时间分配

2005年9月至2006年9月,采用瞬时扫描取样法,在弄岗国家级自然保护区对野生熊猴(Macaca assamensis)进行了连续跟踪观察,收集相关的活动节律和活动时间分配的数据,通过分析其与气候及食物组成的关系,从中探讨影响熊猴活动节律及活动时间分配的因素。研究结果发现:在熊猴的日活动节律中,觅食活动表现为逐渐增强的趋势,最高峰值出现在下午15:00,休息活动在中午出现一个小的高峰;其日活动节律表现出明显的季节性差异,主要表现为旱季大部分时间段的觅食强度均高于雨季,且早上7:00-10:00出现一段长时间的休息,而雨季里长时间的休息则发生在中午12:00-14:00;在活动时间分配上,熊猴平均花费39.6%时间用于休息,33.2%用于移动,18.3%用于摄食,5.1%用于理毛,2.4%用于玩耍,1.4%用于其他行为,其活动时间分配也表现出明显的季节性差异;与雨季相比,在旱季熊猴明显增加用于觅食的时间,而相应地减少用于休息的时间,活动时间分配的季节性变化明显与食物组成的变化有关;在不同年龄组个体间,青少年猴花费更多的时间用于玩耍,而用于休息和理毛的时间明显少于成年猴。     

Seasonal Variation in the Activity Patterns and Time Budgets of <Emphasis Type="Italic">Trachypithecus francoisi</Emphasis> in the Nonggang Nature Reserve,China

Activity patterns and time budgets are 2 important aspects of animal behavior that researchers use to investigate ecological influences on individual behavior. We collected data on activity patterns and time budgets in 1 group of François’ langurs (Trachypithecus francoisi) from August 2003 to July 2004 in the Nonggang Nature Reserve, Guangxi Province, China, via instantaneous scan sampling method with 15-min intervals. The diurnal activity pattern of François’ langurs showed morning and afternoon feeding peaks, with a midday resting peak. Seasonal change was apparent in the activity pattern: 2 significant feeding peaks occurred in the dry season and only 1 significant feeding peak in the rainy season. The group spent an average of 51.5% of the daytime resting. Feeding and moving accounted on average for 23.1% and 17.3% of the activity budget, respectively. Subjects spent little time on social activities, averaging 2% for grooming and 5.5% for playing. Their time budgets showed significant seasonal variation: they spent a greater proportion of time on feeding and less time on resting and grooming in the dry season than in the rainy season. They also differed among different sex-age classes: immatures spent more time playing, whereas adults devoted more time to resting, feeding, and grooming. Correlations between time budgets and food items or food availability clearly indicated that François’ langurs might adopt an energy-maximizing strategy when preferred foods were scarce in the dry season.     

环境和社会因素对喀斯特石山猕猴日活动节律和活动时间分配的影响

2007年9月至2008年8月,以广西弄岗国家级自然保护区内的一大群野生猕猴为研究对象进行连续的跟踪观察,采用瞬时扫描取样法收集其食物组成、活动节律和活动时间分配的数据,旨在探讨影响石山猕猴活动节律及活动时间分配的环境和社会因素。结果表明：研究猴群的日活动节律表现为上午和下午的觅食高峰以及中午的休息高峰,这与白天温度和日照强度的变化有关。日活动时间分配中,休息42.1%,移动34.5%,觅食15.1%,玩耍2.0%,理毛5.2%,其他行为1.3%。猴群依据食物的变化调整活动时间分配,其中用于移动的时间与食物中果实的比例成显著的正相关性。不同性别年龄组个体的活动时间分配表现出显著差异,主要表现为成年雄性花费更多的时间休息,而成年雌性花费更多的时间觅食和社会性理毛,未成年个体则花费更多的时间玩耍。与同域分布的小群猕猴相比,本研究群花费更多的时间休息,较少的时间觅食。这些差异可能与盗食农作物有关。     

秦岭川金丝猴的昼间活动时间分配

1999年12月至2004年11月,通过野外直接跟踪观察,采用焦点取样和瞬时记录的方法,对秦岭川金丝猴西梁猴群的昼间活动时间分配进行了研究。将川金丝猴的昼间活动定义为摄食、休息、理毛、移动和其他行为5 种类型,结果发现：川金丝猴昼间活动中有两个摄食高峰,分别在每天的上午和下午,中午有一个较长时间的休息期。川金丝猴昼间的主要行为类型是摄食和休息（约占全部活动时间的62%）,各种行为依所占时间比例依次为：摄食（34.64 ± 1.70%）、休息（26.83 ± 1.69%）、理毛（11.58 ± 0.96%）、移动（18.93 ±1.08%）和其他行为（8.02 ± 0.82%）。在不同季节各种行为类型所占时间比例有明显的变化,表现为摄食和休息行为占昼间活动时间比例在夏（75.00%）、秋季（74.15%） 大于春（52.74%）、冬季（56.64%）;而移动、理毛和其他行为所占时间比例之和在夏（25.00%）、秋季（ 25.85%） 则明显小于冬（ 43.4%）、春季（47.3%）。不同季节间的活动时间分配经非参数检验（ Kruskal Wallis H test,N = 199）,理毛行为（ x² =11.438,df = 3,P < 0.05）有显著差异,摄食（x² = 20.469,df = 3,P < 0.01）、休息（x² = 27.235,df = 3,P < 0.01）、移动（x² = 24.110,df = 3,P < 0.01） 和其他行为（ x² = 60.810,df = 3,P < 0.01） 季节间差异非常显著;除摄食行为外,其余各行为类型的时间分配在不同性别-年龄组间也存在显著差异（P < 0.05）。     

三种笼养灵长类活动时间分配的比较研究

采用焦点动物法和连续记录法对3种笼养灵长类活动时间分配进行比较研究。结果表明,川金丝猴Rhi-nopithecus roxellanae和熊猴Macaca assamensis用于休息的时间多于猕猴Macaca mulatta,移动时间则相反;川金丝猴和猕猴用于理毛的时间多于熊猴。不同性别年龄组之间的活动时间分配有差异,主要表现在川金丝猴母亲用于理毛的时间明显多于成年儿子,用于玩耍的时间则相反;猕猴成年雄性的移动时间和理毛时间多于成年雌性,玩耍时间则少于成年雌性;熊猴成年雌性的移动时间多于成年雄性,与成年个体相比,幼体花更多的时间用于玩耍,而相应的减少了其休息时间。     

Activities and feeding behavior of free-ranging pregnant baboons

The activities and feeding behavior of pregnant baboons, Papio cynocephalus,were studied in two free-ranging groups in Amboseli National Park, Kenya. Both the environmental conditions and the age of their fetuses influenced females’ activity budgets and feeding behavior. Females fed more and rested less during the dry season than during the wet season, and most females spent progressively more time feeding and less time grooming than expected (based upon the amount of recent rainfall) as their fetuses matured. During the wet season, females also devoted significantly greater proportions of feeding time to consumption of grass blades, leaves, and flowers and significantly smaller proportions of their feeding time to feeding upon seeds, corms, and fruit. As their fetuses became older, the proportion of time spent feeding upon seeds increased, and the proportion of time spent feeding upon grass blades and fruit declined.     

The Effect of Climatic Factors on the Activity Budgets of Barbary Macaques (Macaca sylvanus)

Climatic conditions can significantly affect the behavior of animals and constrain their activity or geographic distribution. Barbary macaques (Macaca sylvanus) are one of the few primates that live outside the tropics. Here we analyze if and how the activity budgets of Barbary macaques are affected by climatic variables, i.e., air temperature, relative humidity, rainfall, and snow coverage. We collected scan sampling data on the activity budgets of four groups of macaques living in the Middle Atlas Mountains of Morocco from June 2008 to January 2011. This habitat is characterized by extreme seasonal changes, from cold and snowy winters to hot and dry summers. The activity budgets of the macaques differed across months but not across the time of day (with the exception of time spent feeding). The monkeys spent significantly more time feeding or foraging when there was no snow than when snow coverage was moderate or major. Daily rainfall was positively related to resting time and negatively to time spent moving or in social behavior. Air temperature was negatively related to time spent feeding or foraging. Finally, time spent on social behavior was significantly lower when relative humidity was high. These data indicate that environmental factors significantly affect the time budgets of endangered Barbary macaques, a species that has been little studied in the wild. Our findings support previous studies on temperate primates in showing that snow coverage can have negative consequences on the feeding ecology and survival of these species.     

白头叶猴日活动时间分配及其季节性变化

<正>活动节律和活动时间分配是动物行为学研究的两个重要方面,它们直接与动物的代谢和能量收支相关,但它们又会随环境不同不断调整。通过比较不同生态条件下动物的活动节律和活动时间分配,     

Diurnal activity budget of African buffalo (Syncerus caffer Sparrman, 1779) in Chebera Churchura National Park,Ethiopia

One of the fundamental questions in animal ecology concerns the activity pattern of animals and the environmental and intrinsic factors that influence such dynamics. This study tested the hypotheses that activity time budgets of the African buffalo appeared to vary by season and times of day and predicted that buffalo would express unequal proportion of time for different activity patterns during the wet and dry seasons in Chebera Churchura National Park (CCNP). An investigation on the diurnal activity budget of the African buffalo was carried out during the wet and dry seasons of 2012–2014 in the Chebera Churchura National Park, Ethiopia, using focal‐animal sampling method. Buffalo spent a greater proportion of the time in feeding and resting/ruminating activities in both the wet and dry seasons. Feeding and resting (lying down and standing) were the predominant activities (87.14% of the diurnal active period), 48.95% time spent feeding during the dry season and 44.91% during the wet season. There was a significant decrease in feeding and an increase in resting from dry seasons to wet seasons. Daytime grazing and resting periods during the wet season were estimated to be 5.39 h and 4.98 h, respectively. Morning and the late afternoon activity peaks were more pronounced during the dry season than the wet season. Therefore, feeding and resting time was influenced by the time of day and the seasons. But there were no significant difference in time allocation for other activities in both the wet and dry seasons. The study has implications for understanding animal activity budget across species, particularly relationships between temperature and season.     

Variation in Spatial Cohesiveness in a Group of Japanese Macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

The spatial cohesiveness of a group is an important element that characterizes the social structure of group-living species. Moreover, remaining cohesive is crucial if individuals are to coordinate their activities and reach collective decisions. We measured interindividual spacing in a group of wild Japanese macaques (Macaca fuscata) to assess the spatial cohesiveness of a social group quantitatively. We used simultaneous focal animal sampling, with 2 observers recording individuals’ locations with a global positioning system (GPS) during 3 seasons. Interindividual distances differed among seasons; they were short in autumn (mean ± SD: 25.6 ± 20.1 m), intermediate in winter (mean ± SD: 46.3 ± 35.7 m), and long in summer (mean ± SD: 62.3 ± 47.1 m). Measurements taken in summer revealed extremely wide spacing (maximum: 1225 m), suggesting subgrouping. Distances also varied with activity during each season; they were short during resting and grooming, intermediate during foraging, and long during moving. Group cohesion was also influenced by food distribution. More group members were ≤20 m of the focal individual during foraging on clumped food than foraging on scattered food in each season, and the group foraged on clumped food most frequently in autumn. Individuals were also likely to aggregate at resting/grooming sites and clumped food patches and to disperse when moving within a day. These results demonstrate that Japanese macaques show considerable variation in spatial cohesiveness both within short time periods, e.g., 1 d, and among seasons, and that they adjust group cohesiveness flexibly depending on the food conditions and foraging tactics.     

圈养阿拉伯狒狒日活动时间格局及季节差异

动物个体行为的生态效应主要反映在它们的活动节律和时间分配上.2012年11月至2013年10月,在杭州野生动物园,用瞬时扫描取样法对1群圈养阿拉伯狒狒的日活动节律和活动时间分配进行了调查研究.结果表明: 1)在活动时间分配中,主要活动(频次百分率≥5%)包括休息(42.5%)、游动(16.2%)、玩耍(10.0%)、摄食(9.8%)、理毛(9.4%)和观望(7.3%);频次百分率低于5%的称为次要活动,共4.8%.2)阿拉伯狒狒的活动时间分配具有明显的季节性变化,冬季摄食和观望时间增加,玩耍和理毛时间减少,但休息和游动行为没有显著季节差异.3)阿拉伯狒狒通常在上午和下午分别出现摄食高峰,并随着气温逐渐转冷,摄食活动逐渐增加,出现3个摄食高峰;这可能是动物在寒冷季节为抵御寒冷而采取的补充能量的一种策略.4)在温暖季节,阿拉伯狒狒只有中午一个休息高峰和一个理毛高峰,更多时间用于游动和玩耍,表现为多峰特征;在寒冷的冬季,休息行为也体现了多峰特征,但其游动、玩耍和张望时间节律并不与温暖季节有明显差异.因此,杭州地区圈养阿拉伯狒狒的冬季日活动节律与许多灵长类(如黑冠长臂猿、维氏冕狐猴和绒毛猴等)不一样,采取摄食增加能量,而非休息减少能量消耗的策略.总之,阿拉伯狒狒的行为体现出明显的节律性,且其活动节律具有季节性差异,这些可能是阿拉伯狒狒适应于圈养环境特有的食物资源和温度变化的影响而形成的.     

贺兰山岩羊不同年龄和性别昼间时间分配的季节差异

2003年11月-2004年10月在宁夏贺兰山国家级自然保护区苏峪口国家森林公园,在每个月的11-20日采用瞬时扫描取样法,利用SJ-1型电子事件记录器观察记录岩羊的取食、休息、运动、站立和其他行为.冬季每天的观察时间为800-1700,春、秋季为700-1800,夏季为600-2000.采用非参数估计中的2个相关样本的Wilcoxon检验、2个独立样本的Mann-Whitney U检验和K个独立样本的Kruskal-Wallis H检验对岩羊不同性别、年龄以及季节变化进行了比较.结果表明,①岩羊行为的年时间分配,取食行为最高[(63.49±7.82)%];其次是休息[(19.32±6.79)%];运动和站立较低[分别占(8.03±0.91)%和(6.66±0.57)%];其他行为所占比例最低[(2.67±0.34)%];一年中7月取食行为最低,休息最高;而在2月取食最高,运动、站立和其他行为变化不大.②春夏秋3个季节的昼间取食行为均呈现2个(清晨和黄昏)明显高峰;冬季1天中取食均达到60%以上,其他行为明显高于另外3个季节.③不同年龄年取食行为成体和亚成体>幼体;休息成体和亚成体<幼体;运动和站立幼体<成体和亚成体;亚成体的运动高于成体,而站立低于成体.④不同性别月取食行为雄性在2月最高,其余月份均低于70%;而雌性虽然也是2月最高,但从11月-次年2月均超过70%;休息行为雌雄接近;运动行为雄性高于雌性.贺兰山岩羊的昼间时间分配与许多温带有蹄类动物一样,食物数量和质量的变化以及岩羊自身不同的生长阶段和生理时期是决定其昼间时间分配的主要因素.     

Habitat Quality and Activity Budgets of White-Headed Langurs in Fusui,China

Within a species, habitat quality may be a factor causing different activity budgets between populations. The habitat of white-headed langurs (Trachypithecus leucocephalus) has been seriously disturbed in Fusui Rare and Precious Animal Nature Reserve, China, where we carried out a study of their socioecology from September 1997 to September 1998. We collected data on langur activity budgets from the main population located in the central part of a group of limestone hills. We classified habitat quality into 4 grades according to the extent of human disturbance. We showed that the two main study groups of white-headed langurs spent on average 50% of time resting, 13% feeding, 18% moving (including foraging), 11% grooming, and 7% playing. Langur time budgets showed no significant seasonal change, but they differed among different sex-age classes. Infants and juveniles spent about 20.3% of time playing, whereas adults spent only 0.2% playing. The group in high quality habitat engaged less in feeding and more in playing than the group in low quality habitat did. Habitat quality influenced the playing time of young white-headed langurs and may be vital to their successful maturation.     

Effects of seasonal and climate variations on calves’ thermal comfort and behaviour

The aim of this study was to measure the effect of season and climate variations on thermal comfort and behaviour of 6-month-old dairy calves housed in a semi-opened shelter to develop animal-based indicators for assessing animal thermal comfort. The ultimate purpose was to further exploit the use of those indicators to prevent thermal stress by providing appropriate care to the animals. Measurements were taken for winter and summer seasons. Results showed that season significantly influenced (P?≤?0.01) the lying down behaviour of calves by reducing the time spent lying, from 679.9 min in winter to 554.1 min in summer. Moreover, season had a significant influence (P?≤?0.01) on feeding behaviour. In detail, the total length of feeding periods was shorter in winter, 442.1 min in comparison to 543.5 min in summer. Time spent drinking increased significantly (P?≤?0.001), from 11.9 min in winter to 26.9 min in summer. Furthermore, season had a significant influence (P?≤?0.001) on self grooming behaviour which was 5.5 times longer in duration in winter than in summer (1,336 s vs 244 s). It was concluded that calves’ thermal comfort is affected by seasonal and climate variations and that this can be assessed by measuring behaviour with animal-based indicators, such as lying down, resting, standing up, feeding, rumination, drinking and self grooming. The indicators developed may be a useful tool to prevent animal thermal stress by providing appropriate housing and handling to calves under seasonal and climate challenge.     

Seasonal Effects on Reconciliation in Macaca fuscata yakui

Dietary composition may have profound effects on the activity budgets, level of food competition, and social behavior of a species. Similarly, in seasonally breeding species, the mating season is a period in which competition for mating partners increases, affecting amicable social interactions among group members. We analyzed the importance of the mating season and of seasonal variations in dietary composition and food competition on reconciliation in wild female Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Yakushima macaques are appropriate subjects because they are seasonal breeders and their dietary composition significantly changes among the seasons. Though large differences occurred between the summer months and the winter and early spring months in activity budgets and the consumption of the main food sources, i.e., fruits, seeds, and leaves, the level of food competition and conciliatory tendency remained unaffected. Conversely, conciliatory tendency is significantly lower during the mating season than in the nonmating season. Moreover, conciliatory tendency is lower when 1 or both female opponents is in estrous than when they are not. Thus the mating season has profound effects on reconciliation, whereas seasonal changes in activity budgets and dietary composition do not. The detrimental effects of the mating season on female social relationships and reconciliation may be due to the importance of female competition for access to male partners in multimale, multifemale societies.     

Sleeping Cave Selection,Activity Pattern and Time Budget of White-Headed Langurs

We describe the activity patterns and time budgets of white-headed langurs that were confined to about 4 km² of Longlin habitat in Fusui County, Guangxi Province, China. Between February and December 1996, we observed 6 langur groups monthly via group focal sampling and continuously recording the behavior of a focal group. Our results indicate that the langur groups selected stone caves on cliffs as sleeping sites. The daily activity pattern outside caves had 8 stages: (1) leaving the cave in the early morning; (2) moving and resting; (3) morning feeding; (4) moving a long distance; (5) resting at noon; (6) afternoon feeding; (7) moving back to the cave, and (8) entering the cave. Over the year, langurs spent a daily average of about 11.5 h outside caves and about 12.5 h inside caves. Moving accounted for 7% (spring), 7% (summer-autumn) and 13% (winter) of the time budget, and langurs spent 9% (summer-autumn) and 14% (spring) to 20% (winter) of their time feeding. Resting accounted for 79% of the time budget in spring, 84% in summer-autumn, and 57% in winter. Sunbathing only occurred in winter and accounted for about 10% of the time budget. One-way ANOVA and multiple range tests demonstrated that time budgets differed significantly among seasons. Langurs spent significantly more time feeding and moving in winter than in spring and summer-autumn, but significantly less time resting in winter than in spring and summer-autumn.     

Enclosure environment affects the activity budgets of captive Japanese macaques (Macaca fuscata)

Individuals adapt to changes in their environment, such as food availability and temperature, by adjusting the amount of time spent in different behavioral activities. These adjustments in behavior should vary across age-sex class according to specific physiological and social needs. We studied the activity budgets of three social Japanese macaque groups inhabiting either vegetated or nonvegetated enclosures in order to compare the effects of access with vegetation, as both food and substrate on resting, feeding, grooming and moving activities over a 12-month period. Daily access to natural foods for monkeys in the vegetated enclosure seems to be largely responsible for the differences in daily time budgets of these three groups. Resting time in all three groups was longer than the time devoted to other activities. Resting and moving time in the two nonvegetated enclosures was significantly longer than in the vegetated enclosure. In contrast, feeding and grooming time was significantly longer in the vegetated enclosure. Seasonal variation in time spent feeding, resting and grooming was significantly effected by enclosure type. In all three enclosures, immatures, particularly females, spent more time feeding and moving, whereas adults spent more time resting. Significant monthly variation in time spent by age-sex class was noted only for feeding and resting. Interestingly, in the vegetated enclosure, time spent feeding on natural vegetation was equal to the amount of time spent feeding on provisioned food. This suggests that factors other than energetic and nutritional needs may be important determinants of the activity budget of the species. These results have important implications for the enrichment of captive primates and our understanding of the maintenance of activity patterns by primates in the wild.     

白马雪山自然保护区响古箐滇金丝猴活动时间分配

2008 年6 月至2009 年5 月,在白马雪山国家级自然保护区南端的响古箐(北纬27°37′,东经99°22′),采用瞬时扫描取样法对一群数量约480 只的滇金丝猴群的活动时间分配进行了研究。研究期间共观察取样1 609 h,扫描个体数为260 546。研究表明,响古箐滇金丝猴用38.8% 的时间取食、27. 4% 的时间移动、20. 9%的时间休息、12.9% 的时间从事其他类型活动。滇金丝猴活动时间分配季节性差异明显。冬季,滇金丝猴的取食时间长于其他季节,达到41. 5% ;秋季,取食时间在全年最短(36.5% )。夏季的高温没有影响滇金丝猴的移动,它们移动的时间达到32.8% ,高于其他3 个季节。寒冷的冬季,响古箐滇金丝猴为了减少能量损失,用于休息的时间达到24.4% 。秋季,滇金丝猴从事其他类型活动的时间(15.7% )多于另外3 个季节。研究发现,
绝对移动时间与昼长和月平均温度呈明显正相关(P < 0. 01),绝对休息时间与月平均温度呈负相关(r =- 0. 585,P = 0.046),但降水量的多少与猴群各种活动时间分配之间没有明显的相关性(P > 0.05)。食物贡献率与滇金丝猴日活动时间分配也存在相关性。取食松萝比例与猴群移动时间呈负相关(r = - 0.902,P <0.001),与休息时间呈正相关(r = 0.860,P < 0.001)。猴群取食树叶比例增加时,它们移动时间也随之增加(r = 0.832,P = 0.001)。夏季,滇金丝猴取食竹笋的比例增加时,猴群移动时间也明显增多。以上结果表明,滇金丝猴活动时间分配受食物资源状况、昼长季节性变化和气候条件的影响。     

Time budget of the green monkey,Cercopithecus sabaeus: Some optimal strategies

Members of a single group of green monkeys spent, on average, 44.8% of their waking time foraging, 46.7% resting, and 8.5% in social activities, over 1 year. There was significant variation in activity budgets over months (ranging from 35 to 55% of the time spent feeding). Diurnal rhythms of feeding and ranging were influenced by the daily cycle of temperature in predictable ways in different seasons: in the dry season, activity was reduced if it was too hot or too cold, while the temperature in the wet season did not affect activities. Feeding was also synchronized among individuals on a finer time scale, irrespective of the time of day. There was closer synchrony when feeding on less common foods. An ecological model of foraging time and energetics was tested, using estimates of the costs and benefits of foraging and predicting how these are optimally balanced in relation to the food density. Both feeding time and distance traveled increased as food availability increased. Costs and benefits were balanced over several days. Comparisons between populations of Cercopithecus aethiopswere made; differences in time budgets were compared with differences in the availability and quality of food. Insufficient comparative data are available for firm conclusions about the role of different energetic and nutritive strategies in population differences.     

Time budgets of Nuthatches Sitta europaea with supplementary food

Individual time budgets during winter and spring were studied in a population of Nuthatches Sitta europaea in south-central Sweden. In late winter birds spent about 90% of daytime foraging, males feeding somewhat less than females, possibly due to male dominance. In early spring, foraging time decreased to about 65% of daytime activity, and sex-related differences became more pronounced. Males then increased time spent resting and singing. Females were nestbuilding, spent less time resting and also devoted more time to foraging than did males. Prior to the observation periods, food density for part of the population was experimentally increased by supplying sunflower seed. In spite of this, time budgets in the experimental group were similar to those of the control group. It is possible that the intensity of foraging, rather than actual time devoted to it, was influenced by food density and that, as a result, birds in the experimental group were able to maintain a higher level of vigilance. Also, an increase in food supply may be less important to a hoarding bird such as the Nuthatch. However, the result remains that Nuthatch time budgets in winter and spring are not readily influenced by an increase in food density.