Search patterns, resource regeneration, and ambush locations impact the competition between active and ambush predators.

Many predators ambush prey rather than pursue them or shift between foraging modes. Active predators typically encounter prey more frequently than ambush predators. I designed a simulation model to examine whether this always holds and how active and ambush predators fare in capturing mobile prey. Prey foraged for clumped resources using area-restricted search, shifting from directional movement before resource encounter to less directional movement afterward. While active predators succeeded more than ambush predators, the advantage of active predators diminished when ambush predators were positioned inside resource patches rather than outside. I investigated the impact of eight treatments and their interactions. For example, regeneration of prey resources increased the difference between ambush predators inside and outside patches, and uncertain prey capture by predators decreased this difference. Several interactions resulted in outcomes different from each factor in isolation. For instance, reducing the directionality level of active predators impacted moderately when applied alone, but when combined with resource regeneration it led to the worst success of active predators against ambush predators inside patches. Ambush predators may not always be inferior to active predators, and one should consider the key traits of the studied system to predict the relative success of these two foraging modes.

Metabolomics analysis of larval secretions reveals a caste-driven nutritional shift in a social wasp colony.

Social wasps exhibit a unique nutritional cycle in which adults feed larvae with prey, and larvae provide adults with larval secretions (LS). LS serves as a vital nutritional source for adults, contributing to the colony's health and reproductive success. The LS nutrient composition has been previously reported in various wasp species, yet these analyses focused solely on worker-destined larvae, overlooking the potential caste designation effects on LS composition. Using metabolomics techniques, we analysed and compared the metabolite and nutrient composition in LS of queen- and worker-destined larvae of the Oriental hornet. We found that queen-destined LS (QLS) contain greater amounts of most metabolites, including amino acids, and smaller amounts of sugars compared to worker-destined LS (WLS). The amino acid-to-sugar ratio in QLS was approximately tenfold higher than in WLS. Thus, as the colony transitions from the production of workers to the production of reproductives, it gradually experiences a nutritional shift that may influence the behaviour and physiology of the adult nest population. This caste-specific metabolite profile and nutrient composition of LS reflect the differences in the diet and physiological requirements of worker- and queen-destined larvae and may play a critical role in caste determination in social wasps.

Flour beetles prefer corners over walls and are slowed down with increasing habitat complexity.

Movement affects all key behaviours in which animals engage, including dispersal and habitat use. The red flour beetle, known as a cosmopolitan pest of stored products, was the subject of our study. We examined whether the beetles preferred corners, walls or open areas, and how turns or obstacles in corridors delayed the beetles' arrival at a target cell. Beetles spent significantly more time in corners than expected by chance, while they spent considerably less time in open areas than expected. However, no significant difference was observed between areas with two or three surrounding walls. This could be attributed to the beetles' stronger attraction to corners than crevices or the insufficient proximity of the third wall to the other two. Movement through the corridor was delayed by turns or obstacles, expressed in arrival probabilities, arrival times, time in the corridor or movement speed. Obstacles on the corridor's perimeter had a stronger effect on the beetle movement than those in the corridor's centre owing to the beetles' tendency to follow walls. The research is important also for applied purposes, such as better understanding beetle movement, how to delay their arrival to new patches, and where to place traps.

Experimental arena settings might lead to misinterpretation of movement properties.

Movement is an important animal behavior contributing to reproduction and survival. Animal movement is often examined in arenas or enclosures under laboratory conditions. We used the red flour beetle (Tribolium castaneum) to examine here the effect of the arena size, shape, number of barriers, access to the arena's center, and illumination on six movement properties. We demonstrate great differences among arenas. For example, the beetles moved over longer distances in clear arenas than in obstructed ones. Movement along the arena's perimeter was greater in smaller arenas than in larger ones. Movement was more directional in round arenas than in rectangular ones. In general, the beetles stopped moving closer to the perimeter and closer to corners (in the square and rectangular arenas) than expected by chance. In some cases, the arena properties interacted with the beetle sex to affect several movement properties. All these suggest that arena properties might also interact with experimental manipulations to affect the outcome of studies and lead to results specific to the arena used. In other words, instead of examining animal movement, we in fact examine the animal interaction with the arena structure. Caution is therefore advised in interpreting the results of studies on movement in arenas under laboratory conditions and we recommend paying attention also to barriers or obstacles in field experiments. For instance, movement along the arena's perimeter is often interpreted as centrophobism or thigmotaxis but the results here show that such movement is arena dependent.

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.

Behavioral differences between pit-building antlions and wormlions suggest limits to convergent evolution.

Antlions and wormlions are distantly related insect taxa, both digging pits in loose soil and ambushing arthropod prey. Their hunting method, which is rare in the animal kingdom, is a clear example of convergent evolution. There is little research directly comparing the 2 pit-building taxa. Using the same experimental platform to investigate how they respond to biotic and abiotic environmental factors enables an examination of their convergence and its limits. We investigated the response of antlions and wormlions to 3 factors common in their daily life: disturbance to the pits, prey arrival, and conspecific competitors. Although both increased the pit size following disturbance, wormlions increased it faster than antlions. Antlions responded to prey faster than wormlions, but wormlions improved their response time over days. The most diverging response was toward conspecifics. Whereas antlions relocated their pits fast in response to increasing conspecific density, wormlions never relocated. We suggest explanations for the behavioral differences between the taxa. Our results imply that despite the similar hunting method of the 2 taxa they may differ greatly in their behavior, which in turn might have consequences for their habitat use and population dynamics.

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.

Egg-Laying Behavior of Cataglyphis niger Ants Is Influenced More Strongly by Temperature Than Daylength.

Temperature and photoperiod are the two most important factors that affect all aspects of animal life. We conducted two experiments to examine the effect of temperature and photoperiod on egg laying and development in the desert ant Cataglyphis niger. In the first experiment, we examined the effect of decreasing temperatures and shortening daylength on egg-laying behavior. An additional treatment was exposure to natural autumn conditions. Decreasing temperatures impaired egg laying much more than shortening daylength. The effect, however, was rapidly reversible when raising the temperature. When the outdoor treatment was brought inside the lab at a suitable temperature, queens started laying eggs as well. In the second experiment, we first kept the colonies under warmer temperatures and moved them gradually to cooler temperatures, 1-20 days after the eggs were laid. The probability of eggs developing into larvae and pupae under cooler temperatures was positively influenced by the exposure duration to warmer temperatures before the temperature switch. When the eggs developed into larvae, longer exposure to warmer temperatures before the temperature switch led to faster development. However, when the eggs disappeared (and were probably eaten), longer exposure to warmer temperatures before the temperature switch led to slower egg disappearance. We suggest that the decision to lay eggs is reversible to some extent because the workers can consume the eggs if conditions deteriorate. We suggest that this reversibility reduces the cost of laying eggs at the wrong time.

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.

A guide to area-restricted search: a foundational foraging behaviour.

Area-restricted search is the capacity to change search effort adaptively in response to resource encounters or expectations, from directional exploration (global, extensive search) to focused exploitation (local, intensive search). This search pattern is used by numerous organisms, from worms and insects to humans, to find various targets, such as food, mates, nests, and other resources. Area-restricted search has been studied for at least 80 years by ecologists, and more recently in the neurological and psychological literature. In general, the conditions promoting this search pattern are: (1) clustered resources; (2) active search (e.g. not a sit-and-wait predator); (3) searcher memory for recent target encounters or expectations; and (4) searcher ignorance about the exact location of targets. Because area-restricted search adapts to resource encounters, the search can be performed at multiple spatial scales. Models and experiments have demonstrated that area-restricted search is superior to alternative search patterns that do not involve a memory of the exact location of the target, such as correlated random walks or Lévy walks/flights. Area-restricted search is triggered by sensory cues whereas concentrated search in the absence of sensory cues is associated with other forms of foraging. Some neural underpinnings of area-restricted search are probably shared across metazoans, suggesting a shared ancestry and a shared solution to a common ecological problem of finding clustered resources. Area-restricted search is also apparent in other domains, such as memory and visual search in humans, which may indicate an exaptation from spatial search to other forms of search. Here, we review these various aspects of area-restricted search, as well as how to identify it, and point to open questions.

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.

Negative effects of fluctuating temperatures around the optimal temperature on reproduction and survival of the red flour beetle.

Whereas the vast majority of animals in nature experience daily or seasonal thermal fluctuations, most laboratory experiments use constant temperatures. We examined the effect of fluctuating temperatures on reproduction and survival under starvation, two important components of fitness. We used the red flour beetle as a model organism, which is a significant pest in grain mills around the world. Fluctuations around the optimal temperature were always negative for the adult survival under starvation. The effect of thermal fluctuations on the number of offspring reaching adulthood was negative as well but increased with the extent of exposure. It was the strongest when the adult parents were kept and the offspring were raised under fluctuating temperatures. However, the later the offspring were exposed to fluctuations during their development, the weaker the effect of fluctuating temperatures was. Moreover, raising the parents under fluctuating temperatures but keeping them after pupation at constant temperatures fully alleviated the negative effects of fluctuations on the offspring. Finally, we demonstrate that keeping the parents a few days under fluctuating temperatures is required to induce negative effects on the number of offspring reaching adulthood. Our study disentangles between the effects of thermal fluctuations experienced during the parental and offspring stage thus contributing to the ongoing research of insects under fluctuating temperatures.

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.

Operant conditioning in antlion larvae and its impairment following exposure to elevated temperatures.

Although ambush predators were previously considered limited in their cognitive abilities compared to their widely foraging relatives, there is accumulating evidence it does not hold true. Pit-building antlions are already known to associate vibrations in the sand with the arrival of prey. We used a T-maze and successfully trained antlions to turn right or left against their initial turning bias, leading to a suitable substrate for digging traps. We present here the first evidence for operant conditioning and T-maze solving in antlions. Furthermore, we show that exposure of second instar larvae to an elevated temperature led to impaired retention of what was learned in a T-maze when tested after moulting into the third instar, compared to larvae raised under a more benign temperature. We suggest that climate change, involving an increase in mean temperatures as well as rare events (e.g., heatwaves) might negatively affect the retention of operant conditioning in antlions, alongside known, more frequently studied effects, such as changes in body size and distribution.

Evidence That Artificial Light at Night Induces Structure-Specific Changes in Brain Plasticity in a Diurnal Bird.

We recently reported that artificial light at night (ALAN), at ecologically relevant intensities (1.5, 5 lux), increases cell proliferation in the ventricular zone and recruitment of new neurons in several forebrain regions of female zebra finches (Taeniopygia guttata), along with a decrease of total neuronal densities in some of these regions (indicating possible neuronal death). In the present study, we exposed male zebra finches to the same ALAN intensities, treated them with 5'-bromo-2'-deoxyuridine, quantified cell proliferation and neuronal recruitment in several forebrain regions, and compared them to controls that were kept under dark nights. ALAN increased cell proliferation in the ventricular zone, similar to our previous findings in females. We also found, for the first time, that ALAN increased new neuronal recruitment in HVC and Area X, which are part of the song system in the brain and are male-specific. In other brain regions, such as the medial striatum, nidopallium caudale, and hippocampus, we recorded an increased neuronal recruitment only in the medial striatum (unlike our previous findings in females), and relative to the controls this increase was less prominent than in females. Moreover, the effect of ALAN duration on total neuronal densities in the studied regions varied between the sexes, supporting the suggestion that males are more resilient to ALAN than females. Suppression of nocturnal melatonin levels after ALAN exhibited a light intensity-dependent decrease in males in contrast to females, another indication that males might be less affected by ALAN. Taken together, our study emphasizes the importance of studying both sexes when considering ALAN effects on brain plasticity.

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.