Genetically-based variation between two spider populations in foraging behavior

Summary Optimal foraging theory is based on the assumption that at least some aspects of foraging behavior are genetically determined (Pyke et al. 1977; Kamil and Sargent 1980; Pyke 1984). Nonetheless, very few studies have examined the role of genetics in foraging behavior. Here, we report on geographical differences in the foraging behavior of a spider (Agelenopsis aperta) and investigate whether these differences are genetically determined. Field studies were conducted on two different populations of A. aperta: one residing in a desert riparian habitat, and the other in a desert grassland habitat. Data from the spiders' natural encounters with prey demonstrated that grassland spiders exhibited a higher frequency of attack than riparian spiders towards 13 of 15 prey types, including crickets and ants. Grassland spiders also had shorter latencies to attack 12 of 15 prey types, including crickets and ants, than riparian spiders. Subsequently, we reared grassland and riparian spiders under controlled conditions in the laboratory and observed their interactions with prey to determine whether the populational differences we found in the field could be genetic. Again, grassland spiders showed a shorter latency to attack prey (crickets, ants) than riparian spiders. These latencies were not significantly affected by the hunger state or age of the spiders. Finally, we reared a second generation (F2) of grassland and riparian spiders in the laboratory and observed their interactions with prey to determine whether the populational differences in the previous generation were due to genetic effects or maternal effects. As before, grassland spiders exhibited a shorter latency to attack prey (crickets) than riparian spiders. We conclude that the foraging differences we observed between these two populations of A. aperta are genetically determined. These differences probably have resulted from either natural selection acting directly on attack frequency and the latency to attack prey, or natural selection acting on traits which are genetically correlated with these aspects of foraging behavior.     

Tubes and foraging behavior in larval Chironomidae: implications for predator avoidance

Summary In laboratory experiments, I studied differential susceptibility of four co-occurring species of chironomids to a predatory damselfly. The chironomids differed in foraging behavior and could be ranked according to the amount of time they spent outside of their tubes. In choice experiments, the predator consistently selected the prey which spent more time out of the tube, and time out of tube was a significant predictor of the predation rate coefficient. Electivity indices, calculated from field samples and diet analyses of the predator, supported the laboratory results. The data suggest that exposure to predators in a heterogeneous prey community is largely determined by tubedwelling behavior.     

Infection of tomato by Tomato Yellow Leaf Curl Virus alters the foraging behavior and parasitism of the parasitoid Encarsia formosa on Bemisia tabaci

Encarsia formosa Gahan is a solitary endoparasitoid that is commercially reared and released for augmentative biological control of whiteflies including Bemisia tabaci (Gennadius). Bemisia tabaci biotypes B and Q are two most invasive species that greatly reduce crop yields in China by feeding on plant sap and by transmitting Tomato Yellow Leaf Curl Virus (TYLCV). The effects of TYLCV infection of tomato on E. formosa foraging on B. tabaci B and Q are unknown. In Y-tube olfactometer assays in the present study, E. formosa significantly preferred TYLCV-infected tomato plants over TYLCV-free plants. The wasp females also significantly preferred TYLCV-infected tomato plants infested with 3rd-instar nymphs of B. tabaci biotype Q over TYLCV-free plants with biotype Q nymphs. However, no significant differences were observed when B. tabaci biotype B was infested on tomato plants. The oviposition bioassays confirmed that TYLCV infection on tomato plants resulted in the recruitment of parasitoids. These results indicate that TYLCV-infection of tomato increase the foraging of E. formosa on B. tabaci, as differs on the B and Q biotypes.     

Specialized predation on plataspid heteropterans in a coccinellid beetle: adaptive behavior and responses of prey attended or not by ants

Two plataspid hemipteran species proliferated on Bridelia micrantha(Euphorbiaceae). Colonies of Libyaspis sp., never attendedby ants, developed on branches, while Caternaultiella rugosa lived at the base of the trunks, mostly in association withCamponotus brutus that attends them in carton shelters. Bothplataspid species are prey of the coccinellid beetle Anisolemniatetrasticta, whose larvae always detected them by contact.When attacked the Libyaspis nymphs cowered, so that the hypertrophiedlateral sides of their tergits made contact with the substrate,but the ladybirds slid their long forelegs under these nymphs,lifted them, and bit them on the ventral face. The Caternaultiellanymphs, which do not have hypertrophied extremities of thetergits, tried to escape at contact with the ladybirds, butwere rarely successful. To capture them, the ladybirds eitheradopted the previous behavior or directly grasped then bitthem. We noted a graded aggressiveness in the ants toward theladybirds according to the situation: no aggressiveness onthe tree branches; stopping the ladybirds that approached theshelters where the ants attended Caternaultiella; and fullattack of ladybirds that tried to capture Caternaultiella nymphssituated outside shelters. The latter behavior can emit analarm pheromone that triggers the dispersion of their congenerswhile attracting attending C. brutus workers. Naive workersare not attracted, so we deduce that this behavior is the resultof a kind of learning.     

长距离迁徙鸟类对应于能量积累状态的取食行为调整

迁徙鸟类能够预计到迁徙过程中对能量需求的增加和迁徙途中获得能量的不确定性。最佳迁徙理论指出：迁徙停留期的一系列决策受到体内能量状态、取食机会和迁徙时间的影响。利用刚完成春季跨越墨西哥湾迁徙的鸫类,我们研究了取食行为、体内能量状态和能量积累速度的相互关系。我们用雾网捕获了停留的鸫,然后给每只鸟进行了环志和称重,并估测了表皮下积累的脂肪。为了检测体内能量积累大小对取食行为的影响,我们把在野外观察到的鸟分成肥、瘦两组。当一天里被捕获鸟的平均体重低于相关种的瘦体重时,这一天被观察到的取食鸟被归到瘦组,反之就属于肥组。我们同时对一部分鸟用有色环进行了标记,以便能在野外观察到它们时能准确地知道每一只鸟的初始能量积累状态。在春天完成跨越墨西哥湾迁徙以后的鸫大约有50％在到达停留地时就已经消耗了所有的表皮下脂肪。与肥组鸟相比,瘦组的个体在停留期扩展了取食方法和取食基底,增加了取食速度。与此相关的是,瘦鸟表现出体重积累更多、速度更快。我们的数据表明迁徙鸟在到达迁徙中途停留地时的能量状态会影响到它们停留期间的取食行为和继续迁徙时的生理机能,从而影响停留期的长短和在迁徙途中停留与否的决定[动物学报51(1)：12—23,2005]。     

A human-induced landscape of fear influences foraging behavior of brown bears

Animals adapt their foraging behavior to variations in food availability and predation risk. In Sweden, brown bears (Ursus arctos) depend on a nearly continuous intake of berries, especially bilberries (Vaccinium myrtillus) during late summer and early autumn to fatten up prior to hibernation. This overlaps with the bear hunting season that starts on 21 August. Bilberry occurrence varies across space, as does human-induced mortality risk. Here, we hypothesize that brown bears select for areas with a high probability of bilberry occurrence across a boreal forest ecosystem in Sweden (H1), and that human-induced mortality risk reduces bear selection for bilberries (H2). In addition, we hypothesized that bears that survived the hunting season avoided bilberry areas associated with high risk, whereas bears that were later killed selected more strongly for berries and less against risk prior to the hunting season (H3). To evaluate our hypotheses, we used resource selection functions to contrast bear GPS relocation data (N = 35, 2012–2015) and random positions within the bearś home range with generalized linear mixed effect models against two focal variables: a map predicting bilberry occurrence and a map predicting human-induced mortality risk. We found that bears selected for areas with a high probability of bilberry occurrence (supporting H1), but avoided these areas if they were associated with and high risk of hunting mortality (supporting H2). The killed and surviving bears did not differ in their selection for bilberries, but they did differ in their selection against risk (partially supporting H3). Surviving bears strongly avoided high risk areas, whereas killed bears responded less to risk and selected for high-risk areas with a low probability of bilberry occurrence. This suggests that killed bears selected for other food sources than berries in high risk areas, which exposed them to human hunters. We conclude that bears respond to a landscape of fear during the berry season and that different foraging strategies may have a direct impact on individual mortality during the hunting season.     

Wax covers in larvae of two Scymnus species: do they enhance coccinellid larval survival?

We tested the protective function of larval wax covers in the two ladybird beetle species, Scymnus nigrinus and S. interruptus, against cannibalism, predation and ant aggression, and its importance for the distribution of both species in the field. Cannibalism was generally very low and not influenced by the presence or absence of the wax cover, or by larval size. Fourth-instar larvae of three ladybird species, Adalia bipuncata, Exochomus quadripustulatus and Harmonia quadripunctata, consumed Scymnus larvae-which are much smaller-regularly, independent of the presence or absence of waxes. By contrast, first-instar larvae of the three species had generally little success when attacking Scymnus spp. larvae. Wax-covered S. interruptus larvae survived significantly more attacks by the predacious carabid beetle Platynus dorsalis than larvae without wax cover. Wax-covered S. interruptus larvae and S. nigrinus larvae survived attacks by workers of the ant species, Lasius niger and Formica polyctena, respectively, significantly more often than larvae without wax covers. We show that, in the field, Scymnus larvae have higher densites in ant-attended resources than in unattended ones and conclude that both Scymnus species benefit from the ability to feed in ant-attended aphid colonies by a reduced predation risk.     

The bottlenose dolphin Tursiops truncatus foraging around a fish farm:Effects of prey abundance on dolphins'' behavior

The extent to which prey abundance influences both bottlenose dolphin foraging behavior and group size in the presence of human activities has not previously been studied.The primary aim of this study was to identify and quantify how wild bottlenose dolphins respond,individually and as groups,to the relative abundance of prey around a fish farm.Detailed views of dolphins' behavior were obtained by focal following individual animals whilst simultaneously collecting surface and underwater behavioral data.A to...     

The bottlenose dolphin Tursiops truncatus foraging around a fish farm：Effects of prey abundance on dolphins＇ behavior

The extent to which prey abundance influences both bottlenose dolphin foraging behavior and group size in the presence of human activities has not previously been studied.The primary aim of this study was to identify and quantify how wild bottlenose dolphins respond,individually and as groups,to the relative abundance of prey around a fish farm.Detailed views of dolphins＇ behavior were obtained by focal following individual animals whilst simultaneously collecting surface and underwater behavioral data.A total of 2150 dive intervals were analyzed,corresponding to 342 focal samples,lasting over 34 hours.Bottlenose dolphins remained submerged for a mean duration of 46.4 seconds and a maximum of 249 seconds.This study provides the first quantified data on bottlenose dolphin diving behavior in a marine fin-fish farm area.This study＇s results indicate that within a fish farm area used intensively by bottlenose dolphins for feeding,dolphins did not modify dive duration.Additionally,underwater observations confirmed that dolphins find it easier to exploit a concentrated food source and it appears that hunting tactic and not group size plays an important role during feeding activities.Thus,bottlenose dolphins appear capable of modifying their hunting tactics according to the abundance of prey.When top predators display behavioral responses to activities not directed at them,the task of studying all possible effects of human activities can become even more challenging.     

Balancing organization and flexibility in foraging dynamics

Proper pattern organization and reorganization are central problems facing many biological networks which thrive in fluctuating environments. However, in many cases the mechanisms that organize system activity oppose those that support behavioral flexibility. Thus, a balance between pattern organization and pattern flexibility is critically important for overall biological fitness. We study this balance in the foraging strategies of ant colonies exploiting food in dynamic environments. We present discrete time and space simulations of colony activity that uses a pheromone-based recruitment strategy biasing foraging towards a food source. After food relocation, the pheromone must evaporate sufficiently before foraging can shift colony attention to a new food source. The amount of food consumed within the dynamic environment depends non-monotonically on the pheromone evaporation time constant—with maximal consumption occurring at a time constant which balances trail formation and trail flexibility. A deterministic, ‘mean field’ model of pheromone and foragers on trails mimics our colony simulations. This reduced framework captures the essence of the flexibility-organization balance, and relates optimal pheromone evaporation to the timescale of the dynamic environment. We expect that the principles exposed in our study will generalize and motivate novel analysis across a broad range systems biology.     

Pea aphid dropping behavior diminishes foraging efficiency of a predatory ladybeetle

Antipredator defensive behaviors are a well‐studied and often crucial part of prey life histories, but little has been done to quantify how such behaviors affect natural enemies, their foraging, and their effectiveness as biological control agents. We explored how the generalist predatory coccinellid Harmonia axyridis Pallas (Coleoptera: Coccinellidae) affects the dropping behavior of the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), and in turn, how that defensive behavior affects the foraging efficiency of the predator. Experimental arenas that allowed or prevented pea aphid dropping were compared to determine how dropping influences the foraging of multiple life stages of H. axyridis: second instars, fourth instars, and adults. Dropping reduced predation on aphids by all ladybeetle life stages. Despite older predators inducing more dropping, aphid dropping reduced predation by approximately 40% across all ladybeetle life stages. Aphid dropping and predator consumption of aphids were both correlated with how much the predator moved, which also increased with predator life stage. We suggest that the high rates of dropping induced by H. axyridis and the subsequent decrease in H. axyridis foraging efficiency may partially explain why H. axyridis is less effective at controlling pea aphids than it is at controlling other aphid species that do not drop.     

Larval development and foraging behavior of Erythrodiplax abjecta (Rambur) (Anisoptera: Libellulidae) in captivity

Understanding the life cycle of Neotropical odonate species is essential given our scant knowledge of this region. In this paper, we examine growth ratio between instars, development patterns, and active/passive feeding behavior of the Andean dragonfly Erythrodiplax abjecta (Rambur, 1842). Larvae were obtained from eggs laid by two females in the laboratory and were maintained at 12–34 °C in individual containers until either their emergence or their death. Larvae hatched 26–57 days after laying, and the total development time was determined as being 316 (SD ± 6.6) days, including 13 instars. Larval instars were characterized using six morphometric variables. The foraging behavior was analyzed considering the time of day and the percentage of the background covered by detritus. The growth ratios between successive instars averaged 1.9 for FW pad length, 1.6 for HW pad length and 1.2 for head width, head length, metathoracic leg length, and total length. Neither the active foraging nor the sit-and-wait foraging behavior were affected by either time of the day or the percentage of background covered by detritus. Erythrodiplax abjecta is univoltine and has a slow lifestyle associated with lentic perennial waters, where larval development and growth rates are low. We provide an equation to estimate the E. abjecta larval instars from field specimens.Given that our results were obtained from the eggs laid by only two females, further studies including a larger sample size are needed to validate their generality.     

The phylogeny of termites (Dictyoptera: Isoptera) based on mitochondrial and nuclear markers: Implications for the evolution of the worker and pseudergate castes, and foraging behaviors

A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. All of these results were very stable and supported with high bootstrap and Bremer values. The evolution of worker caste and foraging behavior were discussed according to the phylogenetic hypothesis. Our analyses suggested that both true workers and pseudergates (“false workers”) were the result of at least two different origins. Our data support a traditional hypothesis of foraging behavior, in which the evolutionary transition from a one-piece type to a separate life type occurred through an intermediate behavioral form.     

Effect of sexual dimorphism in bill length on foraging behavior: an experimental analysis of hummingbirds

We examined whether sexual differences in trophic morphology are associated with sexual differences in foraging behavior through two laboratory experiments on rufous hummingbirds (Selasphorus rufus) designed to compare probing abilities (maximum extraction depths) and handling times of sexes at flowers. Bills of female S. rufus are about 10.5% longer than bills of males, and this difference was associated with sexual differences in foraging abilities. Maximum extraction depths of female S. rufus were significantly greater than those of males, and no overlap between the sexes was observed. Moreover, handling times of females were shorter than handling times of males at flowers having longer corollas (15 mm). Thus, because of their longer bills, female S. rufus have the potential to feed from longer flowers than males, and can do so more quickly. We suggest that no single mechanism is responsible for the evolution of sexual dimorphism in bill lengths of hummingbirds, but rather that the dimorphism probably reflects the combined effects of reproductive role division and intersexual food competition, and possibly, sexual selection.     

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。     

The role of antipredator behavior in an experimental community of jumping spiders with intraguild predation

 The role of direct and indirect interactions in intraguild predation (IGP) was investigated in a laboratory study. The study system contained two spider species, Phidippus audax and Phidippus octopunctatus, and the fruit fly, Drosophila melanogaster. P. audax and P. octopunctatus eat D. melanogaster. P. audax (top predators) also eat P. octopunctatus (intermediate predators). Thus, P. audax and P. octopunctatus compete for the shared resource and also interact as predator and prey. Experiments consisted of two treatments: risk-IGP and full-IGP. In the risk-IGP treatments, I examined the effects of trait-mediated indirect effects generated by antipredator behavior of P. octopunctatus on the survival of fruit flies. P. audax chelicerae were waxed so that P. audax could not attack a prey. The result indicated a significant positive indirect effect of P. audax on the survival of D. melanogaster due to the antipredator behavior of P. octopunctatus (a trait-mediated indirect effect). In the full-IGP treatments, P. audax chelicerae were not restricted, so that it could attack prey; this resulted in decreased survival of D. melanogaster. Because of predation of P. audax on P. octopunctatus, even stronger positive interactions occurred between P. audax and D. melanogaster in full-IGP than in risk-IGP. Received: May 7, 2002 / Accepted: June 27, 2002     

The impacts of Harmonia axyridis cues on foraging behavior of Aphidius gifuensis to Myzus persicae

This study investigated the effect of adult associated cues of the multicolored Asian ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) on the foraging behavior of the aphid parasitoid, Aphidius gifuensis Ashmead (Hymenoptera: Aphididae) with three assays: 1) whole aphid leaf-disc with H. axyridis tracks; 2) aphid leaf-disc with only half contaminated with H. axyridis tracks; 3) aphid leaf-discs without H. axyridis tracks. Foraging behavior of A. gifuensis was recorded using the Observer® XT 11 and EthoVsion® XT 12. In addition, functional responses of A. gifuensis in patches with or without H. axyridis cues were also tested. Aphidius gifuensis females preferred oviposition in arenas where no adult tracks of H. axyridis were present. However, no significant difference between functional responses of A. gifuensis foraging in plants with and without H. axyridis cues was detected. Our study suggests that H. axyridis associated cues could influence the foraging behavior and activity of A. gifuensis under laboratory conditions. On the other hand, the foraging efficiency of A. gifuensis was not significantly affected by H. axyridis walking tracks in a single plant system. Aphidius gifuensis might exhibit flexible behavioral responses to predator associated cues. The risk of intraguild interactions and the possibility of mitigating such risks for parasitoids are important components for ultimately determining the compatibility of biological control agents.     

Spot-fixed fin digging behavior in foraging of the benthophagous maiden goby, Pterogobius virgo (Perciformes: Gobiidae)

 Several patterns of feeding behaviors have been documented in benthophagous fishes. The foraging behavior of the maiden goby, Pterogobius virgo, was studied at Kurahashi Island in the Seto Inland Sea, Japan. Pterogobius virgo foraged mostly on polychaetes by volume from among several available prey items by digging in the sandy bottom. The digging behavior comprised swing of only pectoral fins or of both pectoral fins and body. Pectoral fin swing exposed the cryptic prey within the bottom, and fins and body swing exposed the prey and washed the sediment away. The swings were repeatedly and continuously conducted at a site during the daytime, making a pit several centimeters deep in which the fish was located. After the prey was exposed, the fish immediately and rapidly picked up the prey. Polychaetes were abundant prey in the sediment, occurring in the layer 3–5 cm deep from the bottom surface in the study area. In this goby, spot-fixed fin digging, the first documentation of feeding habits in gobies, may be effective for feeding on the most valuable prey, i.e., polychaetes, which may be otherwise unavailable for this fish. Received: April 24, 2001 / Revised: April 26, 2002 / Accepted: May 7, 2002     

Sources of variability in the transition from extensive to intensive search in coccinellid predators (Homoptera: Coccinellidae)

In an environment structured by habitats, prey patches, and prey, predators such as coccinellids have two movement modes. The extensive search and the intensive search which results from prey capture are adopted for patch localization and exploration, respectively. The variability of changes from extensive search to intensive search was studied in larvae of the aphidophagous coccinellidSemiadalia undecimpunctata to find out their possibility of adaptation to a fluctuating environment. The temporal organization of coccinellid movements appears far more complicated than the generally accepted succession of extensive search, feeding, and intensive search. Their paths are characterized by the presence of time intervals devoted to intensive search before feeding, a highly variable path response after prey consumption (larvae may adopt intensive search immediately, later, or never), and the alternation of time periods devoted to either extensive search or intensive search after prey ingestion. This interindividual variability suggests that coccinellids have the ability to adapt to heterogeneity or short-term changes in environmental conditions, particularly in prey distribution. These results are in favor of the use of these predators in biological control programs.