The combined role of visual and olfactory cues in foraging by Cataglyphis ants in laboratory mazes.

Foragers use several senses to locate food, and many animals rely on vision and smell. It is beneficial not to rely on a single sense, which might fail under certain conditions. We examined the contribution of vision and smell to foraging and maze exploration under laboratory conditions using Cataglyphis desert ants as a model. Foraging intensity, measured as the number of workers entering the maze and arriving at the target as well as target arrival time, were greater when food, blue light, or both were offered or presented in contrast to a control. Workers trained to forage for a combined food and light cue elevated their foraging intensity with experience. However, foraging intensity was not higher when using both cues simultaneously than in either one of the two alone. Following training, we split between the two cues and moved either the food or the blue light to the opposite maze corner. This manipulation impaired foraging success by either leading to fewer workers arriving at the target cell (when the light stayed and the food was moved) or to more workers arriving at the opposite target cell, empty of food (when the food stayed and the light was moved). This result indicates that ant workers use both senses when foraging for food and readily associate light with food.

An experimental critique of the population carrying capacity concept.

Carrying capacity has multiple definitions, but nowadays, it is mainly referred to as the maximum number of individuals of a particular species sustained by the environment. We examined whether multiple populations of two flour beetle species grown under controlled laboratory conditions reach similar asymptotic population sizes when provided with similar amounts of food resources. We demonstrate that the variation in the asymptotic population sizes was considerably larger than that of the initial food resources and that the latter had no significant effect on the former. Our results experimentally contribute to past literature criticizing the carrying capacity concept, demonstrating that there is no single carrying capacity even under strict laboratory conditions. Therefore, we should not expect to often find "carrying capacities" in nature, where resources fluctuate over time, and interspecific interactions are ubiquitous. We suggest that the classic meaning of carrying capacity should be revisited or saved chiefly for didactic purposes.

Snakes on a slope: strong anti-gravitactic responses and differential habitat use in the Saharan horned viper (Cerastes cerastes) in the Negev desert.

The way species use their habitat dictates their intra- and interspecific interactions. We studied the effects of the microhabitat type and slope on the movement behaviour of the Saharan horned viper (Cerastes cerastes) in its natural habitat. This viper occurs in sand dunes and moves mostly by sidewinding. Additionally, we studied the microhabitat preference of desert rodents-the vipers' main prey. We placed the vipers on different natural dune slopes and recorded their behaviour. We found a strong anti-gravitactic response: vipers moved more frequently towards the top of the dune than in any other direction, despite a decrease in stride length with increasing slope. The foraging-related behaviour of the vipers was concentrated in the dune semi-stable areas rather than its stable or shifting sand areas. We measured rodent activity by placing seed trays in the dune allowing the rodents to collect seeds. Rodent activity was the highest in the shifting sands, closely followed by the semi-stable microhabitat. These results suggest the vipers use the semi-stable microhabitat mainly for foraging and may use the shifting sand areas as commuting routes between such areas. This study may be of use for conservation efforts of psammophilic species in desert dunes.

The value of spatial experience and group size for ant colonies in direct competition.

Animals often search for food more efficiently with experience. However, the contribution of experience to foraging success under direct competition has rarely been examined. Here we used colonies of an individually foraging desert ant to investigate the value of spatial experience. First, we trained worker groups of equal numbers to solve either a complex or a simple maze. We then tested pairs of both groups against one another in reaching a food reward. This task required solving the same complex maze that one of the groups had been trained in, to determine which group would exploit better the food reward. The worker groups previously trained in the complex mazes reached the food reward faster and more of these workers fed on the food than those trained in simple mazes, but only in the intermediate size group. To determine the relative importance of group size versus spatial experience in exploiting food patches, we then tested smaller trained worker groups against larger untrained ones. The larger groups outcompeted the smaller ones, despite the latter's advantage of spatial experience. The contribution of spatial experience, as found here, appears to be small, and depends on group size: an advantage of a few workers of the untrained group over the trained group negates its benefits.

Leg or antenna injury in Cataglyphis ants impairs survival but does not hinder searching for food.

Injury is common in nature resulting, for example, from fighting, partial predation, or the wear of body parts. Injury is costly, expressed in impaired performance, failure in competition, and a shorter life span. A survey of the literature revealed the frequent occurrence of injury in ants and its various causes. We examined whether leg or antenna injury impacts food-discovery time and reduces the likelihood of reaching food in workers of the desert ant Cataglyphis niger. We examined the search-related consequences of injury in groups of either 4 or 8 workers searching for food in a short arena, a long arena, and a maze. We conducted a small field survey to evaluate the prevalence of injury in the studied population. Finally, we compared the survival rates of injured versus uninjured workers in the laboratory. Injury was common in the field, with almost 9% of the workers collected out of the nest, found to be injured. Injured workers survived shorter than uninjured ones and there was a positive link between injury severity and survival. However, we could not detect an effect of injury on any of the searching-related response variables, neither in the arenas nor in the mazes tested. We suggest that workers that survive such injury are only moderately affected by it.

Foraging behaviour, habitat use and population size of the desert horned viper in the Negev desert.

The desert horned viper occurs in the dunes of the northwestern Negev desert, Israel. We report on a 2 year study on the viper's behaviour and ecology in its natural habitat. We examined whether the vipers moved faster in a vegetation-dense microhabitat versus an open dune area and detected much slower movement in the former. We nevertheless detected no preference of the vipers for any of the dune areas. We suggest that the vipers trade-off the ease of movement on open areas with prey, which is probably more available in areas with denser vegetation. The activity was higher early in the season and the vipers were mostly active right after sunset, with a second smaller activity peak at sunrise, perhaps searching for burrows to spend the day. Fitting this explanation, movement at the track's end was less directional than at its beginning. We found inter-sexual and between-year differences. For example, females were larger than males in the second year of the study but not in the first one and the population seemed to be smaller in the second year of the study than in its first year. The information we provide on this viper may assist its conservation, as sand dunes are threatened habitats in Israel.

Desert Ants Learn to Avoid Pitfall Traps While Foraging.

Central-place foragers, such as social insects or nesting birds, repeatedly use the same routes from and to their nests when foraging for food. Such species forage more efficiently after accumulating experience. We examined, here, a relatively neglected aspect of such an improvement with experience-the avoidance of pitfall traps. Similar pits are built by antlions, which co-occur with the ants, but they also resemble other natural obstacles. We used the desert ant Cataglyphis niger, common in sandy habitats, and allowed it to forage for three successive runs for a food reward. Ant workers discovered food more slowly and in smaller numbers when pits were in their path. Pit presence also led to longer tracks by ants and slower movement. However, with experience, the ants fell into such pits less often and reached the food more quickly. To understand how past conditions affect current behavior, we investigated whether removing or adding pits led to a different result to that with a constant number of pits. Workers adjusted their behavior immediately when conditions changed. The only carryover effect was the longer tracks crossed by workers after pit removal, possibly resulting from the mismatch between the past and current conditions. Finally, the workers were more likely to fall into pits that were closer to the nest than those that were further away. This is a good example of the advantage that ambush predators can derive from ambushing their prey in specific locations.

Evidence for the effect of brief exposure to food, but not learning interference, on maze solving in desert ants.

Theories of forgetting highlight 2 active mechanisms through which animals forget prior knowledge by reciprocal disruption of memories. According to "proactive interference," information learned previously interferes with the acquisition of new information, whereas "retroactive interference" suggests that newly gathered information interferes with already existing information. Our goal was to examine the possible effect of both mechanisms in the desert ant Cataglyphis niger, which does not use pheromone recruitment, when learning spatial information while searching for food in a maze. Our experiment indicated that neither proactive nor retroactive interference took place in this system although this awaits confirmation with individual-level learning assays. Rather, the ants' persistence or readiness to search for food grew with successive runs in the maze. Elevated persistence led to more ant workers arriving at the food when retested a day later, even if the maze was shifted between runs. We support this finding in a second experiment, where ant workers reached the food reward at the maze end in higher numbers after encountering food in the maze entry compared to a treatment, in which food was present only at the maze end. This result suggests that spatial learning and search persistence are 2 parallel behavioral mechanisms, both assisting foraging ants. We suggest that their relative contribution should depend on habitat complexity.

Urban Pit-Building Insects Are Attracted to Walls for Multiple Reasons.

Whereas most animals find urban habitats to be inferior to natural habitats, some "urban specialist" species thrive there. Wormlions present such an example. Common in Mediterranean cities, they cluster in thin layers of loose soil below man-made shelters. Wormlions are fly larvae that dig pit-traps in loose soil and hunt small arthropods. Our first aim was to determine whether wormlion pits accumulate next to walls. Wormlion pits were indeed closer to walls than expected by chance at most of the study sites. We examined possible factors behind this apparent preference, combining field observations and experiments, laboratory work, and theoretical analysis. We examined the effect of soil depth, particle size, shade, and prey abundance. Each factor provided a partial explanation for the wormlions' proximity to walls, but none provided an overall explanation. We developed a spatially explicit simulation model, demonstrating under which conditions wall-adjacent positions are favored. Finally, we created artificial microhabitats, and placed wormlions either in the center or next to the wall. The wormlions in the center moved over longer distances than those next to the wall and did so more in the wall's direction. The abundance of walls may help to explain the success of wormlions in urban habitats.

Do pit-building predators prefer or avoid barriers? Wormlions' preference for walls depends on light conditions.

Ambush site selection by sit-and-wait predators is a complex process, involving biotic and abiotic considerations, which greatly affect hunting success and costs. Wormlions are fly larvae that dig pit-traps in loose soil and hunt the arthropod prey falling into their pits. They are abundant in urban environments, found below buildings that provide cover, and many of their pits are dug adjacent to walls. We examined here under what conditions wormlions prefer to dig their pits next to walls. We analysed our dataset in two ways: frequency comparisons among the different treatment combinations and a simulation null model assuming random movement. While the frequency comparisons suggested that wormlions avoided the walls under some cases, the simulation null model suggested that a combination of shallow sand and strong light in the centre led to an attraction towards the walls, independent of the wormlions' initial location. We suggest that wall attraction results from the certain amount of shade the walls provide. We also demonstrate that shallow sand and strong illumination are unfavourable microhabitats, either leading to more frequent movement or the digging of smaller pits. We locate our results within the broader context of sit-and-wait predators and of animals' attraction to barriers.

Non-spatial information on the presence of food elevates search intensity in ant workers, leading to faster maze solving in a process parallel to spatial learning.

Experience can lead to faster exploitation of food patches through spatial learning or other parallel processes. Past studies have indicated that hungry animals either search more intensively for food or learn better how to detect it. However, fewer studies have examined the contribution of non-spatial information on the presence of food nearby to maze solving, as a parallel process to spatial learning. We exposed Cataglyphis niger ant workers to a food reward and then let them search for food in a maze. The information that food existed nearby, even without spatial information, led to faster maze solving compared to a control group that was not exposed to the food prior to the experiment. Faster solving is probably achieved by a higher number of workers entering the maze, following the information that food is present nearby. In a second experiment, we allowed the ants to make successive searches in the maze, followed by removing them after they had returned to the nest and interacted with their naïve nestmates. This procedure led to a maze-solving time in-between that displayed when removing the workers immediately after they had reached the food and preventing their return to the colony, and that of no removal. The workers that interacted upon returning to the nest might have transferred to naïve workers information, unrelated to spatial learning, that food existed nearby, and driven them to commence searching. Spatial learning, or an increase in the correct movements leading to the food reward relative to those leading to dead-ends, was only evident when the same workers were allowed to search again in the same maze. However, both non-spatial information on the presence of food that elevated search intensity and spatial learning led to faster maze solving.

Wormlions prefer both fine and deep sand but only deep sand leads to better performance.

Wormlions are small fly larvae that dig pits in loose soil to trap their prey. Similar to other trap-building predators, like spiders and antlions, they depend on the habitat structure for successful trap construction and prey catch. We examined whether sites at which wormlions are present differ in sand depth and particle size from nearby sites, at which wormlions are absent. Next, in the laboratory we manipulated both sand depth and type (fine vs. coarse) to determine their joint effect on microhabitat preference, the size of the constructed pit, wormlion movement, and their latency to respond to prey. We expected better performance by wormlions in fine and deep sand, and the sand in wormlions' natural sites to be finer and deeper. However, in only partial agreement with our expectations, wormlion sites featured finer sand but not deeper sand. In the laboratory, wormlions preferred both fine and deep sand, and moved more in shallow and coarse sand, which we interpret as an attempt to relocate away from unfavorable conditions. However, only deep sand led to larger pits being constructed and to a faster response to prey. The preference for fine sand could, therefore, be related to other benefits that sand provides. Finally, body mass was a dominant factor, interacting with the preference for both deep and fine sand: deep over shallow sand was more favored by large wormlions and fine over coarse sand by smaller ones. Our results suggest that several factors should be incorporated when studying microhabitat selection.

The effect of maze complexity on maze-solving time in a desert ant.

One neglected aspect of research on foraging behavior is that of the effect of obstacles that increase habitat complexity on foraging efficiency. Here, we explored how long it takes individually foraging desert ant workers (Cataglyphis niger) to reach a food reward in a maze, and examined whether maze complexity affects maze-solving time (the time elapsed till the first worker reached the food reward). The test mazes differed in their complexity level, or the relative number of correct paths leading to the food reward, vs. wrong paths leading to dead-ends. Maze-solving time steeply increased with maze complexity, but was unaffected by colony size, despite the positive correlation between colony size and the number of workers that searched for food. The number of workers observed feeding on the food reward 10 min after its discovery decreased with complexity level but not colony size. We compared our experimental results to three simulation models, applying different search methods, ranked them according to their fit to the data and found the self-avoiding random search to fit the best. We suggest possible reasons for the model deviations from the observational findings. Our data emphasize the necessity to refer to habitat complexity when studying foraging behavior.

Context-dependent effects of cold stress on behavioral, physiological, and life-history traits of the red flour beetle.

Animals are exposed in nature to a variety of stressors. While stress is generally harmful, mild stress can also be beneficial and contribute to reproduction and survival. We studied the effect of five cold shock events versus a single cold shock and a control group, representing three levels of stress (harsh, mild, and no stress), on behavioral, physiological, and life-history traits of the red flour beetle (Tribolium castaneum, Herbst 1797). Beetles exposed to harsh cold stress were less active than a control group: they moved less and failed more frequently to detect a food patch. Their probability to mate was also lower. Beetle pairs exposed to harsh cold stress frequently failed to reproduce at all, and if reproducing, females laid fewer eggs, which were, as larvae in mid-development, smaller than those in the control group. However, harsh cold stress led to improved female starvation tolerance, probably due to enhanced lipid accumulation. Harsh cold shock also improved tolerance to an additional cold shock compared to the control. Finally, a single cold shock event negatively affected fewer measured response variables than the harsh cold stress, but also enhanced neither starvation tolerance nor tolerance to an additional cold shock. The consequences of a harsher cold stress are thus not solely detrimental but might even enhance survival under stressful conditions. Under benign conditions, nevertheless, harsh stress impedes beetle performance. The harsh stress probably shifted the balance point of the survival-reproduction trade-off, a shift that did not take place following exposure to mild stress.

Comparison of wormlions and their immediate habitat under man-made and natural shelters: suggesting factors making wormlions successful in cities.

Wormlions are fly larvae that construct pit-traps in loose soil and ambush prey that fall into their pits. They occur in high numbers in cities, below any man-made shelter providing protection from direct sunlight, such as a concrete roof with a thin layer of sand at the ground. Their natural habitat is either caves or any natural structure that provides full shade. We characterized a large urban habitat and compared it to two natural habitats, where wormlions occur in caves. Wormlions were abundant in all studied habitats. Our goals were to understand whether wormlions in the urban habitat perform better than in the natural habitats, and to suggest differences between the habitats that may contribute to their success under man-made shelters. Wormlions in the city reached larger size before pupation, and wormlion clusters there were larger. The studied urban habitat contained more concrete and perennial plants, while the natural habitats comprised of more annuals. We suggest that this concrete, covered with a thin layer of sand, leads to large areas suitable for wormlions. Furthermore, ants were more common in the urban habitat than the natural habitats, referring to their relative proportion of all arthropods collected. We suggest that these small ants provide suitable prey for wormlions, especially in the early stages of their development, when wormlions are limited by prey size. This could explain why they reach larger size prior to pupation. Pits were probably larger because they were constructed by larger individuals. In conclusion, we suggest that wormlions present an interesting case of an insect pre-adapted to urban life.

Consistent differences in foraging behavior in 2 sympatric harvester ant species may facilitate coexistence.

The co-occurrence of 2 similar species depends on their ability to occupy different ecological niches. Here, we compared the consistency of different aspects of foraging behavior in 2 co-occurring harvester ant species (Messor ebeninus and Messor arenarius), under field conditions. The 2 species are active concomitantly and display a similar diet, but M. arenarius features smaller colonies, larger workers on average, and a broader range of foraging strategies than M. ebeninus. We characterized the flora in the 2 species' natural habitat, and detected a nesting preference by M. arenarius for more open, vegetation-free microhabitats than those preferred by M. ebeninus. Next, we tested the food preference of foraging colonies by presenting 3 non-native seed types. Messor arenarius was more selective in its food choice. Colonies were then offered 1 type of seeds over 3 days in different spatial arrangements from the nest entrance (e.g., a seed plate close to the nest entrance, a seed plate blocked by an obstacle, or 3 plates placed at increasing distances from the nest entrance). While both species were consistent in their foraging behavior, expressed as seed collection, under different treatments over time, M. ebeninus was more consistent than M. arenarius. These differences between the species may be explained by their different colony size, worker size, and range of foraging strategies, among other factors. We suggest that the differences in foraging, such as in food preference and behavioral consistency while foraging, could contribute to the co-occurrence of these 2 species in a similar habitat.

The interplay between maze complexity, colony size, learning and memory in ants while solving a maze: A test at the colony level.

Central-place foragers need to explore their immediate habitat in order to reach food. We let colonies of the individually foraging desert ant Cataglyphis niger search for a food reward in a maze. We did so for three tests per day over two successive days and an additional test after a time interval of 4-20 days (seven tests in total). We examined whether the colonies reached the food reward faster, consumed more food and changed the number of workers searching over time, within and between days. Colonies' food-discovery time shortened within and between days, indicating that some workers learnt and became more efficient in moving through the maze. Such workers, however, also forgot and deteriorated in their food-discovery time, leveling off back to initial performance after about two weeks. We used mazes of increasing complexity levels, differing in the potential number of wrong turns. The number of workers searching increased with colony size. Food-discovery time also increased with colony size in complex mazes but not in simple ones, perhaps due to the more frequent interactions among workers in large colonies having to move through narrow routes. Finally, the motivation to solve the maze was probably not only the food reward, because food consumption did not change over time.

The Effects of Temperature and Diet during Development, Adulthood, and Mating on Reproduction in the Red Flour Beetle.

The effects of different temperatures and diets experienced during distinct life stages are not necessarily similar. The silver-spoon hypothesis predicts that developing under favorable conditions will always lead to better performing adults under all adult conditions. The environment-matching hypothesis suggests that a match between developmental and adult conditions will lead to the best performing adults. Similar to the latter hypothesis, the beneficial-acclimation hypothesis suggests that either developing or acclimating as adults to the test temperature will improve later performance under such temperature. We disentangled here between the effect of growth, adult, and mating conditions (temperature and diet) on reproduction in the red flour beetle (Tribolium castaneum), in reference to the reproduction success rate, the number of viable offspring produced, and the mean offspring mass 13 days after mating. The most influential stage affecting reproduction differed between the diet and temperature experiments: adult temperature vs. parental growth diet. Generally, a yeast-rich diet or warmer temperature improved reproduction, supporting the silver-spoon hypothesis. However, interactions between life stages made the results more complex, also fitting the environment-matching hypothesis. Warm growth temperature positively affected reproduction success, but only when adults were kept under the same warm temperature. When the parental growth and adult diets matched, the mean offspring mass was greater than in a mismatch between the two. Additionally, a match between warm adult temperature and warm offspring growth temperature led to the largest offspring mass. These findings support the environment-matching hypothesis. Our results provide evidence for all these hypotheses and demonstrate that parental effects and plasticity may be induced by temperature and diet.

The involvement of sand disturbance, cannibalism and intra-guild predation in competitive interactions among pit-building antlion larvae.

Competition in trap-building predators such as antlion larvae is a complex biotic interaction, potentially involving exploitation competition, sand throwing (i.e., interference competition), cannibalism and intra-guild predation. We investigated the short-term behavioral and developmental responses of the strict sit-and-wait antlion predator Myrmeleon hyalinus to sand disturbance (i.e., quantification of the impact of severe sand throwing), and to con- and hetero-specific competition by a larger sit-and-pursue antlion species Lopezus fedtschenkoi. We found that antlions subjected to sand disturbances reduced their pit construction activity and relocated less often. Furthermore, the reduction in pit construction activity was stronger among antlions subjected to disturbances prior to feeding. Almost no death occurred during the sand disturbance experiment, but as expected, disturbances caused reductions in the relative growth rates of antlions. This negative effect was stronger in the group exposed to sand disturbances prior to feeding. The presence of the sit-and-pursue competitor led to reductions both in pit construction and in relocation activities of M. hyalinus. Although the per-capita food supply was identical in both experiments, only 48% of M. hyalinus larvae survived the competition experiment, and this pattern was consistent between the con- and hetero-specific treatments. However, in the presence of hetero-specific competitors, the relative growth rate of surviving larvae was significantly lower than that measured in the presence of con-specific competitors. Our study demonstrates that investigating the different components of complex biotic interactions can markedly improve our understanding of how these different factors interact to influence the behavior and life history of organisms.

A morphological and life history comparison between desert populations of a sit-and-pursue antlion, in reference to a co-occurring pit-building antlion.

Although most antlion species do not construct pits, the vast majority of studies on antlions focused on pit-building species. We report here on a transplant experiment aiming to test for morphological and life history differences between two desert populations of a sit-and-pursue antlion species, Lopezus fedtschenkoi (Neuroptera: Myrmeleontidae), originating from habitats, which mainly differ in plant cover and productivity. We raised the antlion larvae in environmental chambers simulating either hyper-arid or Mediterranean climate. We found significant differences in the morphology and life history of L. fedtschenkoi larvae between the two populations. For example, the larvae originating from the more productive habitat pupated faster and had a higher growth rate. In agreement with the temperature-size rule, antlions reached higher final mass in the colder Mediterranean climate and exhibited a higher growth rate, but there was no difference in their developmental time. Observed differences in morphology between populations as well as those triggered by climate growing conditions could be explained by differences in size allometry. We also provide a quantitative description of the allometric growth axis, based on 12 morphological traits. Comparing the responses of L. fedtschenkoi with those observed in a co-occurring pit-building antlion indicated that there were neither shape differences that are independent of size nor was there a difference in the plasticity level between the two species.