同种或异种干扰对花鼠分散贮藏点选择的影响

同种和异种干扰竞争可以增强花鼠(Tamias sibiricus)分散贮食强度,但对其贮藏点选择的影响还不清楚.在半自然围栏内,研究了同种(干扰源为不同性别花鼠)或异种(干扰源为大林姬鼠,Apodemus peninsulae)干扰竞争对花鼠贮藏点选择的影响.结果发现:(1)同种和异种干扰竞争均显著降低了花鼠在高竞争区内的贮藏比例;(2)同种个体存在时,雌性花鼠显著降低了在高竞争区内的贮藏比例,雄性花鼠则显著增加了在中竞争区的埋藏比例.结果表明,花鼠贮藏食物时会避开高竞争区域以降低同种或异种的盗食损失,同时雌雄个体对同种竞争干扰的响应有所差异.     

Head raising during foraging and pausing during intermittent locomotion as components of antipredator vigilance in chipmunks

Although many studies of vigilance examine head raising in foraging, grooming or resting animals, pauses during intermittent locomotion are rarely considered from the perspective of vigilance, and no studies have compared head raising and pausing in the same system. We videotaped central place foraging chipmunks, Tamias striatus, as they approached a patch, collected sunflower seeds, and left to return to their burrows. There was a strong similarity between head raising during foraging and pausing during intermittent locomotion. Chipmunks paused more frequently when moving towards the patch than when leaving the patch. Chipmunks in the patch raised their heads at an intermediate rate, which tended to decrease with time in the patch. Pauses and the duration of motionless periods during head raises were very short (∼0.4 s), and their frequency distributions were similar. Animals remained motionless during 22% of the time spent approaching the patch, 14% of the time spent in the patch and 7% of the time spent leaving the patch. Rates of pausing and head raising generally decreased with short-term familiarity (number of trips to the patch) and with long-term familiarity (proximity of the patch to the burrow). Trials with higher pause rates when approaching the patch had higher head-raising rates in the patch. Whether the focal individual was solitary, dominant or subordinate in a dyad, or competing with multiple chipmunks in the patch had no effect on pausing or head raising. In a separate experiment, exposure to a model hawk increased pause and head-raising rates. We conclude that head raising during foraging and pausing during locomotion serve a similar vigilance function, that this vigilance is directed towards detection of predators rather than conspecifics, and that time allocated to vigilance is sufficient to significantly reduce foraging rates and affect many space use and foraging decisions.     

被动式电子标签用于花鼠种群动态研究的可行性

在野生动物研究过程中经常需要对动物个体进行标记。传统的标记方法存在着标记物易脱落、可能影响被标记动物的生理和行为、难于进行个体识别等缺点。被动式电子标签(passive integrated transponder, PIT)是基于射频识别技术的电子标签,可用于体内标记,具有不易脱落消失且能够进行个体识别的优点。我们在凉水国家级自然保护区以PIT标签为标记物,采用标志重捕技术开展花鼠(Eutamias sibiricus)的种群动态研究,同时对应用PIT标签的可靠性进行了研究。2011年秋季至2012年春季共标记花鼠个体140只,其中86只被重捕至少1次。2011年秋季标记的72只个体中有38只在2012年春季被重捕。重捕的花鼠个体注射口愈合完好,体重无明显降低。结果表明PIT标签是一种安全可靠、识别迅速、数据准确、不易丢失、可循环使用的体内标记电子标签,适用于基于个体识别的小型哺乳动物生态学研究。     

The placement, recovery, and loss of scatter hoards by eastern chipmunks,Tamias striatus

Although eastern chipmunks typically larder hoard food in theirburrows, some scatter hoarding occurs, especially by newly emergedjuveniles and by females with young. To examine patterns ofscatter hoard placement and their adaptive significance, weused standard food provisioning tests followed by continuousmonitoring to determine the longevity and fate of scatter hoards.Longevity of scatter hoards was low (median 74 min) and pilferagewas high (46%). Chipmunks placed scatter hoards away from thepatch, closer to their burrow and within the defended area ofprimary use. Scatter hoard placement was affected by the numberof competitors at the patch. However, juveniles and femaleswith young differed slightly in their responses. competitorsthat approached scatteihoards were sometimes chased away bythe scatter hoard's owner, and the scatter hoard was moved toanother location by the owner. Females with young recoveredtheir largest scatter hoards first, but there was little additionalpattern in scatter hoard recovery. An experiment using simulatedscatter hoards showed that scatter hoards farther from a patchof food lasted longer and that disturbing and marking scatterhoards reduced their longevity. These observations indicate:(1) that scatter hoard placement is more related to avoidingpilferage of completed caches than to rapidly sequestering fooditems from an ephemeral patch and (2) that optimal cache distributionmay be affected by the cache-defense ability of individuals,by the relationship between competitor density and cache securitynear the patch, and by the territorial behavior of neighboringconspecifics.     

Conspecific Avoidance During Foraging in Venturia canescens (Hymenoptera: Ichneumonidae): The Roles of Host Presence and Conspecific Densities

We explored the effects of the presence of conspecifics on host patch choice decisions made by the parasitoid Venturia canescens. Different odor sources were located in plastic boxes at the end of each arm of a glass Y-tube olfactometer. In a set of experiments, odor sources were either (a) host kairomone patches with or without conspecifics (5 or 20) or (b) two odor sources located in successive boxes (a host kairomone patch and a patch with 20 conspecifics in one arm versus a host patch and an empty patch in the other). Our results indicate that V. canescens avoids competition only at high conspecific densities. Avoidance occurs only when foraging wasps perceive the combined odors from host kairomones and conspecific females. Separating the host patch from conspecific parasitoids does not perturb avoidance behavior.     

The biological function of snake mobbing by Siberian chipmunks: I. Does it function as a signal to other conspecifics?

The signal effect of mobbing of snakes (a signal effect alerts conspecifics of the presence of a predator) by Siberian chipmunkEutamias sibiricus asiaticus was examined. A pair of chipmunks, familiar with each other, were moved from their rearing pen into a test box, where they were kept undisturbed for acclimatization. Then the anterior portion (or head) of an anesthetized Japanese rat snake was exposed in a corner of the box. Several minutes later, one of the chipmunks discovered the snake and began mobbing it. This was followed by the discovery of the snake by the other chipmunk. Although the time span from the introduction of the snake to its discovery by the first chipmunk was quite different between trials, the time elasped from the snake discovery by the first chipmunk to that by the other chipmunk was relatively constant and significantly shorter than the former time span in all trials. These results strongly support the premise that the mobbing has a signal effect.     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

To mark the host or the patch: Decisions of a parasitoid searching for concealed host larvae

We found evidence for patch marking in the parasitic wasp Halticoptera laevigata (Hymenoptera: Pteromalidae) foraging for concealed hosts. Wasps attack larvae of the fruit fly Myoleja lucida (Diptera: Tephritidae) in fruits of honeysuckle. A special feature of this host-parasitoid system is the limited food supply of a patch (i.e. a fruit of honeysuckle), which allows the successful development of only a single host fly larva. Females of the parasitoid H. laevigata were found to mark the host patch with a pheromone and to abandon the patch following oviposition into a single host larva. Field data revealed that eggs of the parasitoid were spread out evenly among infested patches, with several larvae of the host fly left unparasitized in those patches that contained more than one host. Since many parasitic insects mark the parasitized host after oviposition, we assumed host marking to be the ancestral character state and studied the patch-marking behaviour of H. laevigata as a derived character state as an alternative foraging strategy. We used stochastic dynamic modelling to investigate under what conditions mutant (patch) markers would be able to invade a population of normal (larval) markers. The models suggested that, under a variety of conditions, wasps marking the patch obtained higher fitness than wasps only marking the larva. Consequently, the results from our model predict the evolution of the patch-marking behaviour found in the empirical investigation. Finally, we discuss alternative pathways to the evolution of patch marking and point out under what circumstances the evolution of a patch-marking behaviour can generally be expected.     

Discrimination of previously searched,host-free sites byMicroplitis croceipes (Hymenoptera: Braconidae)

The ability of free- ranging parasitoids to discriminate between previously visited and unvisited sites containing host kairomone (caterpillar frass) but not hosts was tested. Females of Microplitis croceipes,a host specialist and plant generalist larval parasitoid of Helicoverpa (Heliothis) zea,were allowed to fly freely in a simulated plant patch in a flight chamber. Wasps spent less time searching frass sites previously searched by themselves or by conspecifics than unsearched frass sites. In addition to chemical marking, spatial memory of visual cues was implicated as a mechanism for discriminating against self-visited, host-free sites.     

Effect of retentive markers on the dynamics of settlement: The case of arthropod silk

During a settlement decision, the presence of conspecifics is crucial to species subject to Allee effects, for which the number of founders affects the subsequent growth of the colony. Marking the area (physically or chemically) conveys information about the number of conspecifics present in a new patch. Here, we study how an individual affinity for the marker affects the dynamics of a foundation process. A generic population model is presented, in which marking and affinity for the marker are at stake. Our results show that population size thresholds can appear, below which settlement is not possible. This model is then used to study the dynamics of migration and aggregation in a set of interconnected populations. We show that affinity for the marker can induce asymmetries in the population distribution. Anelosimus eximius is a social spider subject to Allee effects, for which silk potentially acts as a marker. We test our predictions with field experiments involving two populations of A. eximius in a Y-shaped setup. The agreement between our experimental and theoretical results strongly supports the validity of the model. This allows us to use the model to estimate a realistic set of parameters of biological significance to this social spider.     

The Reproductive State of Female Voles Affects their Scent Marking Behavior and the Responses of Male Conspecifics to Such Marks

During the breeding season, the reproductive condition of female mammals changes. Females may or may not be sexually receptive. We conducted a series of experiments to determine whether reproductive condition of female meadow voles affects their scent marking behavior as well as the scent marking behavior of male conspecifics. In expt 1, females in postpartum estrus (PPE females) deposited more scent marks than females that were neither pregnant nor lactating (REF females) or ovariectomized females (OVX females). In expt 2, male voles scent marked more and deposited more over‐marks in areas marked by PPE females than by REF and OVX females. In expt 3, PPE females deposited more scent marks and over‐marks in areas marked by males than did females in the other reproductive states. The results of these experiments showed that male and female voles may vary in the number, type and location of scent marks they deposit in areas scented by particular conspecifics.     

A dynamic habitat selection game

A patch selection game is formulated and analyzed. Organisms can forage in one of H patches. Each patch is characterized by the cost of foraging, the density and value of food, the predation risk, and the density of conspecifics. The presence of conspecifics affects the finding and sharing of food, and the predation risk. Optimal foraging theory can be viewed as a "1-person" game against nature in which the optimal patch choice of a specific organism is analyzed assuming that the number of conspecifics in other patches is fixed. In the general game theoretic approach, the behavior of conspecifics is included in the determination of the distinguished organism's strategy. An iterative algorithm is used to compute the solution of the "n-person" game or dynamic ESS, which differs from the optimal foraging theory solution. Experiments to test the proposed theory using rodents and seed trays are briefly discussed.     

Quantifying the effects of distance and conspecifics on colonization: experiments and models using the loosestrife leaf beetle, Galerucella calmariensis

The ability of an insect to disperse to new habitat patches is difficult to quantify, but key to the establishment and persistence of populations. In this study, we examined dispersal of the phytophagous chrysomelid beetle, Galerucella calmariensis, which is currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), an aggressive wetland weed. We used a mark, release, and recapture approach to determine how rates of colonization of host patches by this beetle are influenced by the distance of the patch from the source of dispersers, and by the presence of conspecifics at the patch. We released color-coded beetles at six distances from a long, linear patch of purple loosestrife that was divided into segments with and without conspecifics. We observed initial flight directions as beetles left the release points and collected all beetles that settled at the target patch. We found a bias in initial flight toward the target for distances up to 50 m. Over the 7 days of the experiment, beetles arrived at the target from all release points, including the farthest release point, 847 m away. G. calmariensis was strongly attracted to conspecifics when settling after dispersal; 86% of the 582 recovered beetles came from the segments inhabited by conspecifics. The probability of an individual arriving at the patch declined steeply with release distance. This relationship fits a model in which beetles move in a random direction and stop if they intercept the target patch, and where beetles are lost at a constant rate with distance travelled. The dispersal and patch-colonizing behavior of G. calmariensis is likely to have important consequences for the biological control program against purple loosestrife. Received: 23 January 1996 / Accepted:30 September 1996     

Does the Siberian chipmunk respond to the snake by identifying it?

Siberian chipmunks were presented with snakes and 4 control animals (tortoises, frogs, eels, quails) in a large outdoor pen to examine whether the snakedirected behavior of the chipmunk is a response to identifying the snake or simply an exploratory behavior toward generally strange objects. The animals were presented to chipmunks in 3 manners; tethered, anesthetized, and in perforated opaque boxes and in wire netting boxes. In “tethered” and “anesthetized”, chipmunks responded significantly more intensively to snakes than to other animals. When the animals were presented in the boxes, that is, when chipmunks could perceive only olfactory stimulus from the animals, they also showed the strongest response to snakes. The results suggest that 1) snake-directed behavior of chipmunks is not a general exploratory response toward strange objects but a response to identifying the snakes, and that 2) olfaction is an important cue for the chipmunks to identify snakes.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

Behavioral Responses of Siberian Chipmunks toward Conspecifics Applied Snake Scent

Siberian chipmunks are known to gnaw snake-related objects such as skin, urine, feces and anal sac excretion, and apply the gnawed bits to their body fur repeatedly (SSA behavior). After SSA, chipmunks often rub their bodies on nearby wood stumps and branches. The SSA behavior is surmised to have function of information-spreading on snakes to neighboring chipmunks. In the present study, responses of the other chipmunks towards SSA individuals were experimentally analyzed to examine the possibility of this hypothesis. In the experiments, subject chipmunks were presented with 2 kinds of objects; (1) SSA-performed chipmunks in boxes and non-performed chipmunks in boxes, (2) wood boards to which the scent of SSA-performed chipmunks adhered and wood boards to which the scent of non-performed chipmunks adhered. In both cases of (1) and (2), subject chipmunks nosed significantly longer the scent from SSA-performed chipmunks and that they performed SSA only in the contact with the former.     

The effect of pair bonding in Cabrera vole’s scent marking

The Cabrera vole (Microtus cabrerae) is a rare rodent living in patchy grassy areas of the Iberian Peninsula where unpaired individuals of both sexes use scent marking primarily to increase their mate-finding likelihood. Cabrera voles establish long-term pair bonds with opposite-sex conspecifics constituting a breeding pair, which is expected to reduce the efforts in searching for a new mate. Under such circumstances, scent marking as a strategy to increase mate-finding likelihood became useless. Accordingly, we hypothesise that pair bonded Cabrera voles suppress mate-finding scent marking to reduce energetic costs and predation risk. To test this hypothesis, we compared scent-marking behaviour towards a clean substrate, in both paired and non-paired voles. No differences were found in the scent marks’ type and the amount of marks placed by voles in both conditions. We also analysed the scent-marking behaviour of both sex pair bonded voles when exposed simultaneously to a clean substrate, a substrate pre-marked by males and a substrate pre-marked by females. We found no significant differences in scent-marks (urine-marked area and number of faecal boli) across the three types of substrate types. In accordance with our prediction, these results suggest that pair bonded Cabrera voles did not use scent marking for mate finding, thus providing further support to the existence of a monogamous mating strategy. Furthermore, our results fail to support the use of scent marking for territorial defence purposes.     

Meadow Voles (Microtus pennsylvanicus) and Prairie Voles (M. ochrogaster) Differ in Their Responses to Over-Marks from Opposite- and Same-Sex Conspecifics

Over‐marking occurs when one individual deposits its scent mark on the scent mark of a conspecific. Previous studies have shown that meadow voles (Microtus pennsylvanicus) and prairie voles (M. ochrogaster) that were exposed to an over‐mark of two same‐sex conspecifics, later responded similarly to the top‐scent mark but differed in their response to the bottom‐scent mark. In the present study, we examined the responses of meadow voles and prairie voles to same‐sex and mixed‐sex over‐marks to ascertain whether their responses reflect the different tactics which males and females in promiscuous (meadow voles) and monogamous (prairie voles) species use to attract opposite‐sex conspecifics and to compete with same‐sex conspecifics. Males and females of both species spent more time investigating the mark of the top‐scent donor than that of the bottom‐scent donor of an over‐mark. Meadow voles exposed to a mixed‐sex over‐mark spent more time investigating the mark of the opposite‐sex conspecific independently of whether it was from the top‐ or bottom‐scent donor. In contrast, prairie voles spent more time investigating the mark of the opposite‐sex donor if it was from the top‐scent donor. These results suggest that: (i) over‐marking serves a competitive function; (ii) the scent marks of individuals attract multiple mates in promiscuous species such as the meadow vole; and (iii) the scent marks of individuals establish and maintain pair bonds between familiar opposite‐sex conspecifics in monogamous species such as the prairie vole.     

Patch exploitation strategies of parasitic wasps under intraspecific competition

The optimal residence time of a forager exploiting a resourcepatch in the absence of competitors has been much studied sincethe development of the marginal value theorem. However, severalforagers are frequently observed exploiting the same patch simultaneously,and patch residence time has been surprisingly little studiedin such competitive situations. The few theoretical models developedon this topic predict that foragers should engage in a war ofattrition and stay in the patch longer than when foraging alone.We tested this prediction in Pachycrepoideus vindemmiae (Hymenoptera:Pteromalidae), a solitary parasitoid species in which femalesare known to defend the hosts they are exploiting via intraspecificfighting. By measuring the effect of direct (i.e., presenceof conspecifics) and indirect (i.e., presence of already-parasitizedhosts) competition on patch exploitation strategies, we revealedan apparent polymorphism of strategies. Indeed, in competitivesituations, some members of the population tended to retreatalmost immediately from the patch whereas other members tendedto remain, further exploit the patch, and be more involved ininteractions with conspecifics. The proportion of "retreaters"strongly depended on competition intensity and prior experienceof competition. The forager's physiological state (age) alsoaffects patch exploitation strategy and the intensity of interactionsbetween competing females. Our results highlight the necessityfor further theoretical studies that consider cases where contestsbetween foragers are costly and take into account prior experienceof competition and the forager's physiological state.     

不同生活史阶段长爪沙鼠尿标记行为的遗传和空间距离效应

气味标记普遍存在于哺乳动物类群中,它能向同类传达领地占有、社会等级及繁殖状态等多种信息。2014年4月至10月,我们在内蒙古锡林郭勒正蓝旗对长爪沙鼠 (Meriones unguiculatus) 野外种群进行了标志重捕,采用中立场的方式测定了不同社群间同性别长爪沙鼠的尿标记行为。通过建立洞群样地坐标后用勾股弦方法计算群间距离,并用微卫星DNA分子标记估算个体间的亲缘系数,以探讨野生动物标记行为是否受个体间空间和遗传距离的影响。广义线性混合效应分析显示,雄鼠间的尿标记强度在繁殖期和贮食期受空间距离或遗传距离的影响均不显著。繁殖期雌鼠间的尿标记强度受空间或遗传距离的影响亦不明显,但在贮食期,遗传距离对雌鼠间标记的强度有显著影响,且遗传距离与空间距离对标记强度有明显的交互作用。本研究结果暗示亲缘关系和空间距离对长爪沙鼠的尿标记行为具有一定影响,这与长爪沙鼠的繁殖或贮食行为相适应。