东方田鼠警觉对其功能反应的作用格局

植食性哺乳动物功能反应描述了摄入率与植物可利用性变量的动态关系。动物警觉因占用了处理食物时间即觅食中断时间而延迟与下一口食物相遇,引致摄入率降低,进而对功能反应构型产生影响。在新鲜白三叶叶片构成的各类食物密集斑块上,测定东方田鼠觅食行为,建立功能反应函数模型,检验觅食中断对功能反应的作用格局及机制。结果发现,除小叶片斑块觅食中断时间在觅食活动中所占的比例较低外,在大、中型叶片斑块的觅食中断时间比例均达到15.42%-26.82%;尽管,觅食中断使摄入率降低了33%,但东方田鼠功能反应仍为Ⅱ型功能反应。除东方田鼠采食时间及觅食回合时间随叶片重增大保持相对稳定外,处理时间及觅食中断时间均随叶片重增大呈线性递增趋势;采食时间、处理时间及觅食中断时间随口量的增大呈线性递增趋势;采食率随叶片重和口量增大呈指数递减趋势。研究结果揭示,东方田鼠因警觉引起的觅食中断事件是导致采食率及摄入率降低的主要因子。摄入率测定值与模型预测值的线性回归极显著(P < 0.01),表明,新建立的功能反应模型具有良好的可预测性。推测,东方田鼠因警觉引起采食率及摄入率减小的代价,是以延长觅食时间来补偿的。研究结果充分验证了,在可利用性植物密集斑块,由植物大小调控的动物口量决定其摄入率,且受采食和处理食物竞争及觅食中断的制约,其功能反应为Ⅱ型功能反应的假说。     

东方田鼠家族群对成员个体觅食行为的影响

植食性哺乳动物在分享社群觅食带来好处的同时,是否因个体间的相互干扰而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,测定东方田鼠家族群成员个体在食物斑块上的觅食行为序列过程及行为参数,检验家族群存在对成员个体觅食行为的影响。结果发现,东方田鼠家族群雌、雄成员个体的觅食行为参数均无显著差异。然而,与单只个体相比,家族群觅食尽管能显著地缩短成员个体的觅食决定时间,但却显著地降低了成员个体的摄入率。分析觅食行为参数觅食中断时间发现,相较于单只个体,家族群成员个体间因相互干扰而引起的觅食中断时间的增加,不但增大了收获每口食物的时间,而且导致其摄入率下降。检测家族群成员个体各警觉行为动作参数,发现,成员个体间的相互干扰能引致个体的一般扫视、盯视及嗅闻动作时间比例显著增大,尽管直立扫视和静听动作时间比例减少显著,但并未使个体的觅食中断时间减小。结果充分说明,东方田鼠家族群成员个体间的相互干扰能使个体觅食行为参数发生变异,导致觅食中断时间增加,摄入率降低。     

视野受阻对东方田鼠觅食行为的影响

植食性小型哺乳动物的觅食行为是对特定生境的适应性产物。食物斑块周边植被对动物视野遮挡是否通过作用于其觅食活动中的警觉而影响摄入率。采用新鲜白三叶叶片构建东方田鼠密集均质食物斑块,以牛皮纸模拟食物斑块周边植被遮挡田鼠视野,测定其在食物斑块上的觅食行为序列过程及行为参数,检验食物斑块周边植被高度对东方田鼠摄入率的影响。结果发现,个体在不同程度视野受阻条件下食物摄入率无显著差异。分析觅食行为参数动态发现,在不同视野受阻条件下,个体能通过调整各采食回合内警觉行为动作的发生频次和持续时间,维持觅食回合内总的觅食中断时间的稳定,进而保证进食时间的稳定。东方田鼠在不同程度的视野遮挡条件下均能通过行为变异和优化使摄入率保持稳定。结果亦充分说明,东方田鼠在警觉强度上的变化不能反映觅食中断所带来的食物收益减损的代价,但觅食活动中各警觉动作的持续时间的变异却能够明确指示个体摄入率的动态变化,因此,以觅食活动中警觉引起的觅食中断时间代价为线索,检验其对摄入率的影响,是评价植食性小型哺乳动物在不同生境觅食适应性策略的有效的方法。     

捕食风险对东方田鼠功能反应格局的作用

植食性哺乳动物对食物斑块的选择和利用不仅取决于食物的可利用性,且与觅食环境潜存的各种风险紧密关联。捕食风险是否通过作用于动物觅食活动中的警觉影响其功能反应格局。在新鲜白三叶叶片构成的各类食物密集斑块上,测定东方田鼠觅食行为,建立功能反应模型,检验捕食风险对其功能反应格局的作用。结果发现,捕食风险能显著地延长东方田鼠的觅食决定时间,但其摄入率保持稳定,功能反应构型亦未发生改变,仍为Ⅱ型功能反应;除了对照组个体的采食时间随叶片大小增大无明显变动规律外,处理组个体的采食时间及对照组和处理组个体的处理时间、觅食中断时间均随叶片大小及口量的增大呈线性增高趋势,处理组个体的觅食中断时间明显大于对照组个体的;对照组和处理组个体的采食率均随叶片大小及口量呈非线性渐近递减趋势,但处理组个体的采食率较对照组个体的略有降低。结果揭示,在捕食风险压力下,虽然上述觅食参数变异能潜在地降低摄入率,但个体能通过改变觅食活动中各种警觉行为动作如降低嗅闻和静听监视动作的发生频次,增大视觉监视动作比重,以此缓冲捕食风险压力,维持摄入率。摄入率测定值与模型预测值的线性回归极显著,表明,功能反应模型具有良好的预测性。在可利用植物密集斑块,动物觅食活动中的警觉能缓冲捕食风险压力;动物摄入率是由植物大小调控的口量决定的,且受采食与处理食物竞争及觅食中断的制约;其功能反应仍属Ⅱ型功能反应。     

植食性哺乳动物能量收益增长减速机制的检验

能量收益函数描述了植食性哺乳动物在植物斑块的食物摄入量与在斑块停留时间的函数关系,为觅食生态学理论的重要组成部分。在新鲜白三叶叶片构成的各类叶片斑块上,测定东方田鼠的觅食行为,建立其能量收益函数模型,分析植食性哺乳动物能量收益增长减速的机制。研究结果表明,东方田鼠觅食大叶片时,叶片干物质收益与停留时间呈非线性渐进函数关系,能量收益函数为渐进函数;觅食中、小型叶片时,叶片干物质收益与停留时间呈线性函数关系,能量收益函数为线性函数;没有检测到东方田鼠的能量收益动态呈分段线性函数或S型函数增长。东方田鼠在大、中型叶片斑块觅食时,随停留时间的增大,口量呈线性或指数递减,而处理时间则呈线性或指数递增,采食时间、间隔时间及咀嚼频次保持相对稳定,瞬时摄入率呈减小趋势;东方田鼠在小叶片斑块觅食时,觅食行为参数口量、摄入率、采食时间、间隔时间、处理时间及咀嚼频次均保持相对稳定。研究结果充分验证了,植食性哺乳动物在消费植被过程中,大型可利用性植物减少,受植物大小调控的动物口量减小,处理时间增加,引起瞬时摄入率降低,导致其能量收益增长减速的假说。     

饥饿和食物单宁酸对东方田鼠(Microtus fortis)食物摄入量和觅食行为的影响

实验室条件下,测定饥饿和食物单宁酸对东方田鼠食物摄入量和觅食行为的影响。结果表明,饥饿使实验个体的食物总摄入量增加,食物摄入率及口量大小随饥饿强度的增大而增加,而觅食频次则无显著改变,实验个体每取食回合的觅食时间呈缓慢增加的趋势,与对照组比较,觅食时间差异不显著。东方田鼠优先选择0％单宁酸食物,次为3%单宁酸食物,而对6%单宁酸食物的摄入量最少。在饥饿条件下,东方田鼠食物摄入率的增加主要源于其口量大小,觅食频次和觅摄食时间对食物摄入量增加的贡献不显著。在饥饿条件下,植食性小哺乳动物并未通过延长觅食时间,降低用于防卫、繁殖活动时间来增加食物摄入量,而是通过增加口量大小,提高其食物摄入率来满足其营养需要。验证了饥饿与植物次生化合物共同作用引起田鼠类动物生理的改变,能影响其食物摄入量及觅食行为的假设。     

饥饿和食物单宁酸对东方田鼠(Microtus fortis)食物摄入量和觅食行为的影响

实验室条件下,测定饥饿和食物单宁酸对东方田鼠食物摄入量和觅食行为的影响。结果表明,饥饿使实验个体的食物总摄入量增加,食物摄入率及口量大小随饥饿强度的增大而增加,而觅食频次则无显著改变,实验个体每取食回合的觅食时间呈缓慢增加的趋势,与对照组比较,觅食时间差异不显著。东方田鼠优先选择0%单宁酸食物,次为3%单宁酸食物,而对6%单宁酸食物的摄入量最少。在饥饿条件下,东方田鼠食物摄入率的增加主要源于其口量大小,觅食频次和觅摄食时间对食物摄入量增加的贡献不显著。在饥饿条件下,植食性小哺乳动物并未通过延长觅食时间,降低用于防卫、繁殖活动时间来增加食物摄入量,而是通过增加口量大小,提高其食物摄入率来满足其营养需要。验证了饥饿与植物次生化合物共同作用引起田鼠类动物生理的改变,能影响其食物摄入量及觅食行为的假设。     

植食性哺乳动物能量收益函数模型的预测性及适用性

以东方田鼠喜食的白三叶叶片作为食物,在保持叶片生物量不变的条件下,改变叶片大小,配置东方田鼠觅食的各类食物大小异质斑块,测定东方田鼠觅食的行为。通过比较几种植食性哺乳动物能量收益函数模型的预测性,评价其适用性。结果发现,没有检测到东方田鼠食物摄入量动态呈S型能量收益函数增长。线性函数模型能准确地预测东方田鼠在中、小型食物斑块的停留时间;在大型食物斑块,尽管,分段线性函数及渐进函数均能很好地拟合东方田鼠的食物摄入量动态,但仅分段线性函数模型能准确地预测其停留时间。线性函数模型及分段线性函数模型是在功能反应机制模型-口量模型基础上建立的,反映了调节摄入率动态的机制,为机制性模型。因此,此2种模型是在口量及摄食站尺度上,探讨动物在食物斑块的能量收益动态及停留时间;而渐进函数模型及S型函数模型均为实验性模型,是在斑块尺度上预测动物能量收益动态及停留时间,未能反映动物摄入率动态,故其预测效果较差。由于此4种模型均未考虑动物在食物斑块搜寻食物及非觅食活动如警觉和逃跑等花费的时间,因而,限制了模型的广泛应用。建议,发展新的模型,促进觅食生态学斑块模型理论研究的深入发展。     

东方田鼠社群气味对家族群成员个体觅食行为的影响

东方田鼠家族群成员个体的觅食行为是否因食物斑块存有家族群自身及非亲缘家族群气味而发生变异,进而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,分别配置家族群自身巢垫物及非亲缘家族群巢垫物作为社群气味,测定东方田鼠家族群在食物斑块觅食时,其成员个体觅食行为的序列过程及参数,检验家族群自身气味及非亲缘家族群气味对成员个体觅食行为的影响。结果表明,家族群自身气味能显著地缩短本群成员个体的觅食决定时间,通过减少成员个体的嗅闻及直立扫视动作时间比例、增大一般扫视、盯视及静听动作时间比例,降低觅食中断时间比例,提高其摄入率;而非亲缘家族群气味则能显著地延长家族群成员个体的觅食决定时间,通过增大家族群成员个体的嗅闻和一般扫视动作时间比例、减小直立扫视、盯视及静听动作时间比例,增大觅食中断时间比例,降低其摄入率。结果揭示,熟悉的社群气味会促使觅食活动中的家族群成员个体,在监测环境风险时,将精力更多地用于观察和监听群内其他成员个体的行为及其发出的警报信息,以便在有效规避环境风险的同时减缓个体间因干扰性竞争对觅食活动所造成的不利影响;而陌生的社群气味会迫使成员个体,将精力由依赖群内其他成员个体的行为转向凭借自身直接警觉周围环境。     

棕色田鼠探究行为对同种个体气味源反应格局的分析

通过观察非动情期的棕色田鼠对麻醉的同种个体身体气味源的探究行为 ,包括雄鼠对麻醉雄鼠和雌鼠的嗅闻、雌鼠对麻醉雄鼠和雌鼠的嗅闻 ,分析了棕色田鼠在社会探究中对不同气味源的反应格局。结果表明 ,棕色田鼠对同种个体肛殖区和面部的嗅闻频次及嗅闻持续时间最多 (P <0 0 5 ) ;对背部、腹部、肋部嗅闻的频次和持续时间较少。在对同一部位的嗅闻中 ,在嗅闻频次上 ,雄鼠对雄鼠的嗅闻、雌鼠对雌鼠的嗅闻以及雌鼠对雄鼠的嗅闻都显著多于雄鼠对雌鼠的嗅闻 (P <0 0 5 ) ;在嗅闻时间上 ,性别组合对同一部位的探究行为没有显著影响 ,但气味源与性别组合有交互作用 (P <0 0 5 ) ,影响探究行为。这些结果说明棕色田鼠的肛殖区和面部是重要的气味源 ,而且在对同一气味源的探究中存在性别差异。但在非动情期 ,异性间的探究并不总是高于同性间的探究     

Intraspecific differences in benefits from feeding in mixed-species flocks

The Madagascar Paradise Flycatcher Terpsiphone mutata and Common Newtonia Newtonia brunneicauda frequently form two-species flocks in the deciduous dry forest of western Madagascar. In T. mutata , some males have long tails, while other males and females have short tails. When foraging in mixed flocks, each type of bird captured prey more rapidly than otherwise, but the degree of increase in feeding rate was smaller in long-tailed males. When in mixed flocks, all T. mutata caught prey on leaves in the canopy where N. brunneicauda foraged. Long-tailed males changed feeding habits from sallying when not in mixed flocks, whereas short-tailed birds showed no change of feeding habit. The elongated tails of long-tailed males may have made their foraging less efficient owing to decreased agility in the canopy. N. brunneicauda is monomorphic and often formed groups of three to five individuals. In monospecific flocks, subordinates fed at low rates on branches owing to frequent hostile encounters. When foraging in mixed flocks, however, subordinates foraged among leaves, and their feeding rates increased because the frequency of intraspecific interference decreased greatly. Dominants did not show any difference in feeding pattern with social situation. Thus, heterospecific flocking was more advantageous for subordinates.     

Mating strategies based on foraging ability: an experiment with grasshoppers

Female mate choice and the benefits of this behavior are criticalaspects of Darwinian sexual selection, but they are seldom documentedbecause it is difficult to identify the male trait(s) that femalesmay be seeking. We conducted experiments with grasshoppers (Melanoplussangutnipes: Orthoptera, Acrididae) to examine this behavior.Males that feed more intensively and select a diet mix thatpermits greater food intake (food intake per body mass per time)in laboratory trials were preferentially selected by females.These better foraging males on average provide greater paternalinvestment (greater spermatophore mass) to the female, whichincreases her reproductive rate (eggs produced per body massper time). However, paternal investment may not entirely explainfemale choice of better foraging males, because these maleswere still selected even if they had their food intake restrictedor had been allowed to recently mate, which reduces spermatophoreproduction. Furthermore, males change their mating strategyin response to female choice and the foraging abilities of surroundingmales. Poorer foraging males attempt forcible copulation ratherthan displaying and allowing female choice. A male will facultativelyswitch between these strategies depending on the foraging abilitiesof the surrounding males. While females attempt to reject forciblecopulation, forcible copulation reduces the frequency with whichfemales successfully copulate with better foraging males. Therefore,males that are less "attractive" to females adopt alternativemating strategies to counter female choice which would excludethem from mating.[Behav Ecol 7: 438–444 (1996)]     

Sex differences in giraffe foraging behavior at two spatial scales

We test predictions about differences in the foraging behaviors of male and female giraffes (Giraffa camelopardalis tippelskirchi Matchie) that derive from a hypothesis linking sexual size dimorphism to foraging behavior. This body-size hypothesis predicts that males will exhibit specific behaviors that increase their dry-matter intake rate relative to females. Foraging behavior was examined at two hierarchical levels corresponding to two spatial and temporal scales, within patches and within habitats. Patches are defined as individual trees or shrubs and habitats are defined as collections of patches within plant communities. Males were predicted to increase dry-matter intake rate within patches by taking larger bites, cropping bites more quickly, chewing less, and chewing faster. Within habitats, males were expected to increase intake rate by increasing the proportion of foraging time devoted to food ingestion as opposed to inter-patch travel time and vigilance. The predictions were tested in a free-ranging population of giraffes in Mikumi National Park, Tanzania. Males spent less total time foraging than females but allocated a greater proportion of their foraging time to forage ingestion as opposed to travel between patches. There was no sex difference in rumination time but males spent more time in activities other than foraging and rumination, such as walking. Within patches, males took larger bites than females, but females cropped bites more quickly and chewed faster. Males had longer per-bite handling times than females but had shorter handling times per gram of intake. Within habitats, males had longer average patch residence times but there was no significant sex difference in inter-patch travel times. There was no overall difference between sexes in vigilance while foraging, although there were significant sex by habitat and sex by season interactions. Although not all the predictions were confirmed, overall the results agree qualitatively with the body-size hypothesis. Sex-related differences in foraging behavior led to greater estimated intake rates for males at the within-patch and within-habitat scales. Received: 20 November 1995 / Accepted: 5 November 1996     

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.     

Feeding time strategies of the fringe-toed lizard,Uma inornata,during breeding and non-breeding seasons

Summary I examined the foraging behavior during the breeding and non-breeding seasons, May and July 1986, of the fringe-toed lizard Uma inornata (Iguanidae). During the breeding season males differ from females in their diet and in their foraging time strategy, males exhibiting time minimization and females energy maximization. In May, plant associated foods were selectively eaten. Males concentrated on flowers, a readily available quick energy food, which reduced foraging time and increased time for reproductive activities. Time budgets indicate that males spend over twice as much time in the open and in movement in May than do females. Females at this time restrict their activities to the cover of perennial bushes, and feed primarily on plant foods (flowers and arthropods). Energy maximization appears to be maintained by both sexes in the non-breeding season when food resources diminished to one-half of those in the breeding season. The lizards were less selective in their July feeding habits, broadening their diets to include ground-dwelling arthropods and foliage. Predation by these lizards follows a wait-ambush mode of foraging.