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.

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.

Can rats and ants exchange information between the horizontal and vertical domains?

Since traveling in nature involves encountering various vertical structures, integration of horizontal and vertical spatial information is required. One form of such integration is to use information acquired in one plane for spatial navigation in another plane. Here we tested whether rats and ants that learned a reward location in a horizontal maze could utilize this information when the maze was rotated to a vertical orientation and vice versa. Rats that were trained in a horizontal Y-maze required more time to reach the reward when the maze was vertically rotated, but they were more accurate in choosing the correct arm. In contrast, rats tested in a horizontal maze after being trained in a vertical maze were less accurate but reached the reward faster. Changes after maze rotation were moderate and non-significant in ants, perhaps since the number of ants arriving at the reward increased over trials, diminishing the effect of maze rotation in ants compared to rats. According to the notion that horizontal spatial information is encoded in more detail than vertical information, the slow performance of rats in the vertical domain could be due to a more physically demanding task whereas their accuracy was due to a preceding detailed horizontal encoding. In contrast, rats in the vertical maze could gather less detailed information and therefore were less accurate in subsequent horizontal trials, where the lower energy cost enabled them to swiftly correct wrong choices. Altogether, the present results provide an indication for transferring spatial information between horizontal and vertical dimensions.

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.

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.

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.

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.