内蒙古达赉湖地区蒙原羚昼间行为时间分配

2007年11月～2008年9月,在内蒙古达赉湖地区采用扫描取样法共对404只次蒙原羚Procapra gutturosa成年个体进行了19 385 min (323.1 h) 的昼间行为时间分配观察,将蒙原羚的行为分为采食、站立、移动、卧息、繁殖和其他等6类.结果 表明:蒙原羚晨昏各有一个采食高峰,在12:00前后还会发生1～2次采食小高峰.卧息常发生在采食高峰后,在晨昏所占比例低.单因素方差分析显示蒙原羚春夏秋冬采食、站立、移动、卧息和其他行为的时间分配均存在显著差异,这表明蒙原羚的行为时间分配有较大弹性,从而适应变化的环境.     

矮岩羊夏季活动节律、时间分配和集群行为

采用扫描取样法（每隔10 min进行一次扫描纪录,记录时间为5 min）对36只矮岩羊夏季昼间节律与时间分配进行了观察和研究,将矮岩羊行为划分为取食、移动、站立、卧息和其他行为等5类。结果表明,矮岩羊在夏季有3个取食高峰（6：00－9：00、10：00－11：00和16：00－20：00）;1个卧息高峰（11：00－15：00）。在时间分配上,用于取食的时间最多（30.14±3.32）%,其次是卧息（26.44±3.59）%,站立（24.90±4.27）%,移动（15.73±1.26）%,而用于其他行为的时间,仅占（2.82±1.00）%。通过不同个体的比较,发现成年雄性用于取食和站立的时间最多,幼体用于移动和其他行为的时间最多,成年雌性用于卧息的时间最多。在矮岩羊集群方面,其集群类型有：混合群、母仔群、雄性群、雌性群和独羊,在夏季集群平均大小为（7.81±8.42）只,2~8只的羊群,占总群数的54.05%;不同集群类型的大小差异极显著（χ2=13.197,dƒ=3,P=0.004）,混合群群体最大,而雄性群最小;雌雄比为1∶0.54,雌幼比为1∶0.26,成幼比为1∶0.17。这些都说明矮岩羊这类独特的行为和种群特征是长期适应金沙江干热河谷的结果。     

不同年龄雄性北山羊发情期时间分配特征

前人研究表明,可多次繁殖的反刍物种其雄性个体在发情期采食时间显著减少。目前有两个假说解释这一现象,即能量摄入最大化假说和能量保存假说。为验证雄性北山羊在不同发情阶段所采取的能量保存策略,作者于2014年10-12月在新疆天山中部采用焦点动物取样法采集数据,采用Kruskal-Wallis检验和Spearman秩相关性检验分析数据。研究发现雄性北山羊成体和亚成体发情期采食时间均显著低于发情前期和发情后期,但二者在不同发情阶段卧息时间无显著变化,发情期采食和卧息时间比亦显著降低。发情期发情行为时间显著高于发情前期和发情后期,非发情时间主要用于采食。发情期采食时间和卧息时间都与发情行为时间呈显著的负相关关系。雌性北山羊发情期采食时间亦显著低于发情前期和发情后期,发情期和发情后期卧息时间显著高于发情前期。本研究结果表明,发情期不同年龄阶段雄羊都主要采取能量摄入最大化策略,但同时也具有部分能量保存策略的特征。     

笼养东北虎雌雄行为差异的比较

1998年4月－2001年3月在哈尔滨动物园,采用瞬时扫描取样法和全事件取样法对5只笼养东北虎（3雌2雄）的昼夜行为时间分配及活动规律进行了研究。结果表明：东北虎的睡眠、卧息、运动和其他行为在时间分配上两性存在一定差异,即雌性的睡眠和卧息时间比雄性多,雌性的运动和其他行为时间比雄性少;但摄食行为的时间分配相差不大。两性日活动规律的差异为雄性的睡眠高峰主要在夜晚,卧息较雌性少且相对集中,运动出现和结束的时间较雌性延迟约1h。对两性及不同个体行为时间分配的单因素方差分析发现,不同个体间仅运动存在显著差异（P＜0.05);两性间睡眠、卧息、运动和其他行为均存在显著差异（P＜0.05)。笼养东北虎两性行为的差异可能与其野外的行为习惯有关,而这些行为习惯又与其担当的性别角色有紧密联系。     

不同发情阶段雄性鹅喉羚的时间分配特征

以往研究表明受发情交配行为制约,一些雄性反刍动物在发情期食物摄入量明显降低。已有两个相关假说解释该现象:能量摄入最大化假说和能量保存假说。作者于2009 ～ 2010 年在卡拉麦里山有蹄类自然保护区研究了雄性鹅喉羚不同发情阶段的时间分配。结果表明雄羚发情期采食时间比例明显下降(37. 9% ),低于发情前期(63. 6% )和发情后期(65. 8% );发情期卧息时间比例(6. 0% )与发情后期相近(5. 4% ),明显低于发情前期(23.2% );发情前期至发情后期采食卧息时间比(分别为2. 7、6. 3、12. 1)显著增加;发情期雄羚站立和移动时间比例明显升高,采食行为时间占非发情行为时间主要部分(86. 4% ),且采食行为与发情行为显著相关。相比之下,雌羚不同发情阶段采食行为时间分配比例相似。总之,除必需投入的发情行为外,发情期雄羚最大化其能量摄入;发情行为的投入是导致发情期雄羚食物摄入量下降的主导因子,雄性能量摄入最大化假说更好地解释了发情期鹅喉羚所采取的能量策略。     

竹巴笼矮岩羊昼间行为节律和时间分配

2008年11月中旬至2009年1月中旬和2009年3月至8月,采用瞬时扫描取样法和目标取样法在竹巴笼自然保护区对矮岩羊的昼间行为节律和时间分配进行了研究,结果表明:矮岩羊的昼间行为活动有明显的节律性,活动高峰出现的时间与矮岩羊昼间取食高峰时间基本一致。取食和卧息是最主要的行为方式,分别占总观察频次的53.83%和27.76%,频次较少的是站立和移动,分别占总观察频次的9.93%和8.24%,其它行为所占频次最少,只有0.24%,昼间2个取食高峰(9:00-11:00, 17:00-19:00)和1个卧息高峰(11:00-16:00),呈现出取食-休息-取食的规律。Kruskal-Wallis H 检验表明,季节因素对矮岩羊取食(P <0.05)、卧息(P <0.05)和其它行为(P <0.05)所占时间影响显著,而对移动(P >0.05)和站立(P >0.05)行为影响不显著。各年龄段之间其它行为所占的时间比例差异显著(P<0.05),而取食(P>0.05)、移动(P>0.05)、卧息(P>0.05)和站立(P>0.05)所占时间比例差异不显著;对比分析不同性别矮岩羊昼间行为时间分配差异,雌性的取食、移动和卧息的时间略高于雄性,而站立和其他行为时间低于雄性;Mann-Whitney U 检验对不同性别矮岩羊昼间的各种行为差异进行分析表明:性别因素对矮岩羊昼间时间分配和行为节律产生一定程度的影响,但差异不显著(P>0.05)。     

异地半圈养藏羚卧息行为适应性的初步研究

藏羚Pantholops hodgsoni是青藏高原特有物种。为了探讨藏羚羊对异地环境的行为学适应,对异地圈养藏羚羊的卧息行为进行了不同性别、不同季节间的比较研究。采用全事件记录法和焦点动物取样法,记录和统计了异地圈养藏羚羊冷季和暖季的卧息行为。研究结果表明,在暖季,雌性和雄性藏羚羊在卧息时间及卧息时间占全天活动时间的比例(卧息比例)上没有差异(卧息时间:Z=-1.126,P0.05;卧息比例:Z=-1.357,P0.05);在冷季,雌性和雄性藏羚羊在卧息时间及卧息比例上也没有差异(卧息时间:Z=-1.14,P0.05;卧息比例:Z=-0.307,P0.05)。雄、雌性藏羚羊的卧息时间及卧息比例没有季节差异(雄性卧息时间:Z=-1.465,P0.05;雄性卧息比例:Z=-1.631,P0.05;雌性卧息时间:Z=-1.742,P0.05;雌性卧息比例:Z=-1.852,P0.05)。季节和性别对藏羚羊卧息行为的影响没有交互作用(F=0.057,P0.05)。总之,藏羚羊卧息行为之间既没有性别差异也没有季节差异,与可可西里地区藏羚羊的卧息行为相比,异地圈养藏羚羊的卧息行为已经调整其行为学适应对策。     

不同季节鹅喉羚昼间行为时间分配特征

2007 年11 月至2008 年10 月在新疆卡拉麦里山有蹄类野生动物保护区研究了鹅喉羚的昼间行为时间分配,共观察鹅喉羚有效个体1 917 只,采集行为数据时间319.5 h,将鹅喉羚的昼间行为分为采食、移动、警戒、休息和“其它”5 类。结果表明:春季雌性5 种行为分配时间所占比例分别为67.5%、2.5%、2.7%、25.9%、1.4%,雄性分别为63.5%、3.4%、4.3%、27.5%、1.3%; 夏季,雌性:43.7%、7.7%、5.6%、42.1%、1.0%; 雄性:51.9%、1.7%、3.1%、41.4%、1.9%; 秋季,雌性:66.3%、5.1%、4.3%、23.6%、0.7%;雄性:68.9%、6.0%、4.2%、20.7%、0.3%;冬季,雌性:68.0%、5.0%、7.0%、19.6%、0.4%;雄性:29.6%、20.6%、19.2%、29.3%、1.3%。采用卡方检验、秩和检验、方差分析比较同季节不同性别各自行为时间分配、同季节雌雄间行为时间分配、同性别不同季节的行为时间分配。结果表明:四季中雌雄各自5 种行为的时间分配均存在显著差异(P>0.05);比较两性间行为时间分配发现,在春夏季,雌雄间各行为时间分配无差异(P>0.05),秋季移动差异显著(P < 0. 05),而冬季5 种行为间均差异显著(P<0.05);季节因素对雌雄行为时间分配的影响显著。鹅喉羚以特定的行为时间分配模式适应季节更替或变化的生理周期,如温度、性别、水分状况、繁殖期、哺乳期等,满足能量及营养需求。     

哺乳期圈养雌獐昼夜行为时间分配及活动节律

2014年7—8月,在浙江省富阳市獐养殖场采用瞬时扫描取样法对圈养条件下哺乳期雌獐昼夜行为时间分配及活动节律进行了调查研究。结果表明:1雌獐的日活动行为时间分配依次为卧息(59.01±2.93)%、走动(13.91±2.39)%、采食(10.60±0.74)%、梳理(7.76±1.10)%、反刍(2.84±0.66)%、站立(2.72±1.01)%、警戒(1.53±0.59)%、社会行为(1.26±0.54)%、排遗(0.36±0.22)%。2雌獐的昼间卧息时间极显著低于夜间(P<0.01),梳理时间显著低于夜间(P<0.05),而走动、采食、反刍、警戒和社会行为的时间极显著高于夜间(P<0.01),站立行为差异无统计学意义(P>0.05)。3一天中,雌獐的采食行为和反刍行为呈现双峰型的节律特征,走动、站立和梳理行为出现了3个高峰期,卧息和社会行为则呈多峰型,其中取食高峰后延迟1~2 h为反刍高峰。4卧息行为分别与采食行为和走动行为呈极显著的负相关关系,而采食行为与走动行为之间呈显著的正相关关系。总之,哺乳期圈养雌獐的日活动行为体现出明显的节律性,且大多数行为时间具有明显的昼夜差异。     

不同饲养条件下海南坡鹿幼体春季昼间行为时间分配及活动节律

2003 年2 月至4 月间, 在海南甘什岭自然保护区采用瞬时扫描取样法对10 头海南坡鹿幼体昼间行为时间分配和活动节律进行了初步研究。结果表明: 坡鹿幼体昼间行为时间分配中摄食、卧息和反刍占有较大的比例;人工圈养和半散放条件下摄食、卧息、反刍、站立和运动行为的昼间时间分配差异性显著( P < 0.01), 半散放条件下幼鹿每天用于摄食和运动的时间比人工圈养的多, 而用于卧息、反刍和站立的时间较少, 其余行为时间比较接近; 在昼间活动节律方面, 圈养条件下摄食有3 个高峰期(09:00～10:00, 11:00～12:00, 16:00～18:00) , 卧息有2 个高峰期(10:00～11:00, 13:00～16: 00) ; 半散放条件下没有明显的摄食高峰期, 主要原因是大部分时间花费在食物的搜索和处理上, 而其卧息高峰出现在14:00 以后。     

Feeding or Resting? The Strategy of Rutting Male Alpine Chamois

Optimal foraging theory suggests that animals normally maximize energy intake to optimize their energy balance. However, when efficiency to assimilate energy falls below the level necessary to ascertain basal energetic requirements, they should shift to an energy saving strategy. Males of many ruminant species considerably reduce their food intake during the rut. Nevertheless, they are commonly assumed to maximize energy intake besides their investments in rutting activities. Based on predictions of optimal foraging theory and the specific ruminant digestive physiology, we propose, however, that rutting males in polygynous species with time consuming mating tactics should instead use an energy saving strategy. Particularly, we predict this to be the case in Alpine chamois (Rupicapra rupicapra rupicapra), a highly polygynous mountain ungulate, of which the males defend mating territories during the rut. By combining observational and telemetry data of eight radio‐collared males we constructed individual 24‐h time budgets, and compared the behavior of males before, during and after the rut. Males spent significantly less time feeding during the rut (0.9 h) compared with before (8.5 h) and afterwards (6.4 and 7.5 h, respectively), whereas time spent lying remained more or less unchanged (pre‐rut; 12.7 h, rut; 13.3 h, post‐rut; 12.9 and 13.9 h, respectively). The ratio of time spent feeding to lying dropped from 0.67 in the pre‐rut period to 0.05 in the rutting period. As a result, males allocated on average approx. 90% of their non‐rutting time to lying, and a negative relationship between rutting and lying time emerged. Hence, males seemed to trade lying time against rutting time. We conclude from these results that male Alpine chamois do not maximize their energy intake during the rut, but rather adopt an energy saving strategy to optimize their energy balance.     

Does a temperate ungulate that breeds in summer exhibit rut-induced hypophagia? Analysis of time budgets of male takin (Budorcas taxicolor) in Sichuan,China

Mammals maximize fitness by optimizing time and energy allocation between reproduction and survival. Describing time budgets is a way to understand a species' constraints in energy allocation. We describe a time budget for male takin (Budorcas taxicolor) in Tangjiahe Nature Reserve, China, to better understand rut-induced hypophagia, which is frequently observed in temperate ungulates that breed in autumn or in winter. Observations generally occurred at two elevations (1200-1600m and 2600-3200m), using 20-min focal animal scan sampling from 2007 to 2009. Feeding behaviors accounted for the majority in takin's time budget (61.1%) during daylight hours, relative to the other observed behaviors, such as rest (14.1%), alert behavior (10.2%) and locomotion (6.8%). We found a negative correlation between feeding behavior and rutting behavior during the rutting season. A ratio of feeding time to resting time increased from pre-rut to rut, while resting behavior did not change significantly across seasons. These results suggest the "energy saving" hypothesis could explain reduced foraging in male takin during the rut, but aspects of the species biology suggest that hypotheses for rut-induced hypophagia developed for other temperate ungulates do not apply to takin. We suggest that the unusual summer rutting season of takin releases males from the energy constraints encountered by temperate ungulates that breed in the autumn and has other benefits for offspring survival. Further research should be conducted on ungulates that exhibit rut during the summer and tropical ungulates that might not experience limited food availability following the mating season to improve our understanding on rut-induced hypophagia.     

Rut-Induced Hypophagia in Male Bighorn Sheep and Mountain Goats: Foraging Under Time Budget Constraints

In polygynous ungulates, the rut imposes constraints on male time budgets that generate a trade‐off between maintenance and reproduction, leading to a reduction in time spent foraging. As mating activities can incur substantial somatic costs, males are expected to spend their ‘non‐rutting’ time recovering during the breeding season. If the diminution in time allocated to foraging by males is only a consequence of time budget constraints, males should keep a similar ratio of time spent foraging to lying to that observed in the pre‐rut, leading to an overall reduction of these two activities (the ‘foraging constraint’ hypothesis). Alternatively, if males adopt an energy‐saving strategy, they should limit energy expenditures by reducing foraging but not lying time, as the energy gains of forage intake may not meet the basal energetic requirements, especially in northern and temperate regions (the ‘energy‐saving’ hypothesis). Here, we contrast these two hypotheses by comparing individual daily time budgets of marked adult bighorn sheep rams (Ovis canadensis) and male mountain goats (Oreamnos americanus) during the pre‐rut and the rut. Concordant results for both species support the ‘foraging constraint’ hypothesis, as sexually‐active males reduced time spent foraging and lying from the pre‐rut to the rut because of an increase in time spent in mating‐related activities. Bighorn sheep rams also increased time spent foraging when not engaged in mating tactics, providing further support for a ‘maximisation’ of energy intake in the absence of reproductive opportunities. Because there are also known physiological changes that occur during the rut which may cause appetite suppression, for example to produce metabolic compounds linked with olfactory communication (the ‘scent‐urination’ hypothesis) or to cope with increased burden of parasites (the ‘parasite‐induced anorexia’ hypothesis), further research should aim at simultaneously testing these current hypotheses to better understand rut‐induced hypophagia and its effects on the life histories of male ungulates.     

To Feed or Not to Feed? Testing Different Hypotheses on Rut‐Induced Hypophagia in a Mountain Ungulate

In many ruminant species, males dramatically reduce forage intake during the rut. To date, different hypotheses have been suggested to explain this rut‐induced hypophagia. To assess the predictions of the main hypotheses, we analysed Alpine ibex (Capra ibex) activity budget and compared the behaviour of males and females before, during, and after the rut. Only males spent significantly less time foraging during the rut than outside of it, whereas females allocated a similar proportion of time to foraging before, during, and after the rut. Our results showed that during the rut males also reduced lying time, while the ratio of time spent feeding to time spent lying did not change for males among periods. In conclusion, during the breeding season males maximized energy intake when not actively engaged in mating activities and rut‐induced hypophagia appeared to result from time budget constraints generated by mating‐related activities. Accordingly, the foraging constraint hypothesis seems appropriate to explain this phenomenon in Alpine ibex males.     

圈养獐春夏季昼间行为时间分配及活动节律

2008 年3 ～7 月,在上海市浦东新区华夏公园进行獐重引入试点,采用瞬时扫描取样法和全事件记录法,对圈养条件下24 只獐(12 雌,12 雄)春夏季昼间行为时间分配及活动节律进行了观察。结果表明:休息是春夏季最主要的行为方式,其次是摄食和运动。獐的昼间行为具有明显的节律性,以晨昏活动为主,主要活动时间集中在06:00 ～08:00 和16:00 ～18:00。单因素方差分析(One-way ANOVA analysis)和Mann-Whitney U 检验表明,在时间分配上,春、夏季休息、站立、排遗、梳理和运动行为差异极显著;雌雄间休息、站立、反刍、梳理和警戒行为差异极显著。雌性春、夏季站立、排遗、梳理、运动和社会行为差异极显著,摄食差异显著;雄性春、夏季休息、摄食、梳理、警戒和社会行为差异极显著,站立、反刍、排遗和运动行为差异显著。本研究结果表明重引入獐的行为时间分配和活动节律与野生种群相似,动物福利得到较好满足,对下一步在上海地区的种群复壮和迁地野放等具有重要意义。     

Contrasting Alternative Hypotheses to Explain Rut‐Induced Hypophagia in Territorial Male Chamois

Male ungulates in temperate environments often show a severe reduction in time spent foraging during the mating season. Several hypotheses have been put forward to explain this phenomenon but, so far, no study investigated the proximate mechanisms underlying rut‐induced hypophagia in ungulates using alternative mating tactics (AMTs). Between the pre‐rut and post‐rut of 2011 and 2012, we collected data on activity budgets, parasite burden and androgen levels of territorial and non‐territorial male Alpine chamois Rupicapra r. rupicapra in the Gran Paradiso National Park (Italy). We aimed to investigate whether AMTs showed similar reduction in time spent foraging during the mating period and to test the predictions underlying alternative hypotheses that may explain rut‐induced hypophagia. Only territorial males showed a significant reduction in time spent foraging during the rut; the lack of correlation between proportion of time spent foraging and androgen metabolites or parasite burden did not fully support the physiological and the parasite hypotheses, while the foraging constraint, the energy‐saving and the physical rest hypotheses could not be discounted. Territorial males decreased the time spent lying down from the pre‐rut to the rut, but not their foraging‐to‐lying‐down ratio. During the mating period, we found negative correlations between time spent foraging or lying down and time spent rutting. Our data suggest that territorial males’ behaviour is more consistent with the foraging constraint hypothesis than with the energy‐saving hypothesis previously suggested. Yet, during the rut territorial males did not maximise their foraging time, and the optimisation of their energy balance could rather depend upon feeding on relatively high‐quality plants. This suggestion – possibly named ‘forage quality hypothesis’ – now requires further investigations. This work showed that alternative mating behaviours may underlie different patterns of foraging strategies: we suggest that tests of alternative hypotheses to explain rut‐induced hypophagia within ungulate populations should not ignore the occurrence of AMTs.     

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.