'Dust you shall eat': The complex nutritional and functional considerations underlying a simple diet.

Animals assimilate macronutrients and mineral nutrients in specific quantities and ratios to maximise fitness. To achieve this, animals must ingest different foods that contain the needed nutrients or facilitate the digestion of those nutrients. We explored how these multidimensional considerations affect the desert isopods (Hemilepistus reaumuri) curious food selection, using field and laboratory experiments. Wild isopods consumed three-fold more macronutrient-poor biological soil crust (BSC) than plant litter. Isopods tightly regulated macronutrient and calcium intake, but not phosphorus when eating the two natural foods and when artificial calcium and phosphorus sources substituted the BSC. Despite the equivalent calcium ingestion, isopods performed better when eating BSC compared to artificial foods. Isopods that consumed BSC sterilised by gamma-radiation ate more but grew slower than isopods that ate live BSC, implying that ingested microorganisms facilitate litter digestion. Our work highlights the need to reveal the multifaceted considerations that affect food-selection when exploring trophic-interactions.

Climate dependence of the macrofaunal effect on litter decomposition-A global meta-regression analysis.

Litter decomposition by microorganisms and animals is influenced by climate and has been found to be higher in warm and wet than in cold and dry biomes. We, however, hypothesized that the macrofaunal effect on decomposition should increase with temperature and aridity since larger animals are more tolerant to aridity than smaller organisms. This hypothesis was supported by our global analysis of macrofauna exclusion studies. Macrofauna increased litter mass loss on average by 40%, twofold higher than the highest previous estimation of macrofaunal effect on decomposition. The strongest effect was found in subtropical deserts where faunal decomposition had not been considered important. Our results highlight the need to consider animal size when exploring climate dependence of faunal decomposition, and the disproportionately large role of macrofauna in regulating litter decomposition in warm drylands. This new realization is critical for understanding element cycling in the face of global warming and aridification.

Prudent burrow-site selection in a landscape of fear.

Prey should select safer breeding sites over riskier sites of otherwise similar habitats. This preference, however, may differ between conspecifics of different competitive abilities if the costs of intraspecific competition overpower the benefits of breeding in a safer site. Our goal was to test this hypothesis by exploring the burrow-site selection of different-sized desert isopods (Hemilepistus reaumuri) near and away from a scorpion burrow. We found that larger females are more likely to occupy burrows than smaller females, regardless of whether these burrows were close or away from scorpion burrows. We also found that larger females stayed longer in safer burrows and that smaller females tended to stay longer in riskier sites even in the absence of direct competition, implying a prudent burrow-site selection. We found no association between male size and the tendency to occupy or to spend time in a burrow, regardless of whether these burrows were close or away from scorpion burrows. Our work highlights the need to consider intraspecific competition when exploring how predators regulate prey behaviour.

Multi-elemental consumer-driven nutrient cycling when predators feed on different prey.

Predators play a fundamental role in cycling nutrients through ecosystems, by altering the amount and compositions of waste products and uneaten prey parts available to decomposers. Different prey can vary in their elemental content and the deposition of elements in predator waste can vary depending on which elements are preferentially retained versus eliminated as waste products. We tested how feeding on different prey (caterpillars, cockroaches, crickets, and flies) affected the concentrations of 23 elements in excreta deposited by wolf spider across 2 seasons (spring versus fall). Spider excreta had lower concentrations of carbon and higher concentrations of many other elements (Al, B, Ba, K, Li, P, S, Si, and Sr) compared to prey remains and whole prey carcasses. In addition, elemental concentrations in unconsumed whole prey carcasses and prey remains varied between prey species, while spider excreta had the lowest variation among prey species. Finally, the concentrations of elements deposited differed between seasons, with wolf spiders excreting greater concentrations of Fe, Mg, Mn, Mo, S, and V in the fall. However, in the spring, spiders excreted higher concentrations of Al, B, Ba, Ca, Cd, Cu, K, P, Na, Si, Sr, and Zn. These results highlight that prey identity and environmental variation can determine the role that predators play in regulating the cycling of many elements. A better understanding of these convoluted nutritional interactions is critical to disentangle specific consumer-driven effects on ecosystem function.

Fear of predators alters herbivore regulation of soil microbial community function.

Fear of predation can affect important ecosystem processes by altering the prey traits expression that, in turn, regulates the quantity and quality of nutritional inputs to soil. Here, we aimed to assist in bridging a knowledge gap in this cascading chain of events by exploring how risk of spider predation may affect grasshopper prey performances, and the activity of various microbial extracellular enzymes in the soil. Using a mesocosms field-experiment, we found that grasshoppers threatened by spider predation ate less, grew slower, and had a higher body carbon to nitrogen ratio. Herbivory increased activity of all microbial extracellular enzymes examined, likely due to higher availability of root exudates. Predation risk had no effect on C-acquiring enzymes but decreased activity of P-acquiring enzymes. We found contrasting results regarding the effect of predation on the activity of N-acetyl-glucosaminidase and leucine arylamidase N-acquiring enzymes, suggesting that predation risk may alter the composition of N-inputs to soil. Our work highlighted the importance of soil microbial enzymatic activity as a way to predict how changes in the aboveground food-web dynamics may alter key ecosystem processes like nutritional-cycling.

Lizards exploit the changing optics of developing chromatophore cells to switch defensive colors during ontogeny.

Many animals undergo changes in functional colors during development, requiring the replacement of integument or pigment cells. A classic example of defensive color switching is found in hatchling lizards, which use conspicuous tail colors to deflect predator attacks away from vital organs. These tail colors usually fade to concealing colors during ontogeny. Here, we show that the ontogenetic blue-to-brown tail color change in Acanthodactylus beershebensis lizards results from the changing optical properties of single types of developing chromatophore cells. The blue tail colors of hatchlings are produced by incoherent scattering from premature guanine crystals in underdeveloped iridophore cells. Cryptic tail colors emerge during chromatophore maturation upon reorganization of the guanine crystals into a multilayer reflector concomitantly with pigment deposition in the xanthophores. Ontogenetic changes in adaptive colors can thus arise not via the exchange of different optical systems, but by harnessing the timing of natural chromatophore development. The incoherent scattering blue color here differs from the multilayer interference mechanism used in other blue-tailed lizards, indicating that a similar trait can be generated in at least two ways. This supports a phylogenetic analysis showing that conspicuous tail colors are prevalent in lizards and that they evolved convergently. Our results provide an explanation for why certain lizards lose their defensive colors during ontogeny and yield a hypothesis for the evolution of transiently functional adaptive colors.

In the land of the blind: Exceptional subterranean speciation of cryptic troglobitic spiders of the genus Tegenaria (Araneae: Agelenidae) in Israel.

Caves have long been recognized as a window into the mechanisms of diversification and convergent evolution, due to the unique conditions of isolation and life in the dark. These lead to adaptations and reduce dispersal and gene flow, resulting in high levels of speciation and endemism. The Israeli cave arachnofauna remains poorly known, but likely represents a rich assemblage. In a recent survey, we found troglophilic funnel-web spiders of the genus Tegenaria in 26 caves, present mostly at the cave entrance ecological zone. In addition, we identified at least 14 caves inhabited by troglobitic Tegenaria, which are present mostly in the twilight and dark ecological zones. Ten of the caves, located in the north and center of Israel, are inhabited by both troglophilic and troglobitic Tegenaria. These spiders bear superficial phenotypic similarities but differ in the levels of eye reduction and pigmentation. To test whether these taxa constitute separate species, as well as understand their relationships to epigean counterparts, we conducted a broad geographic sampling of cave-dwelling Tegenaria in Israel and Palestine, using morphological and molecular evidence. Counterintuitively, our results show that the troglobitic Tegenaria we studied are distantly related to the troglophilic Tegenaria found at each of the cave entrances we sampled. Moreover, seven new troglobitic species can be identified based on genetic differences, eye reduction level, and features of the female and male genitalia. Our COI analysis suggest that the Israeli troglobitic Tegenaria species are more closely related to eastern-Mediterranean congeners than to the local sympatric troglophile Tegenaria species, suggesting a complex biogeographic history.

Predation risk regulates prey assortative mating by reducing the expected reproductive value of mates.

Many animals exhibit size assortative mating (SAM), but how predation affects it remains largely unknown. We hypothesized that predation risk may turn prey less choosy, disrupting SAM, or reduce the expected reproductive value of mates, maintaining SAM but with different size ratio between mates. Using a manipulative field experiment, we found that desert isopods under risk of scorpion predation maintained SAM, but that males that choose and fight over females were on average smaller for a given female size. Less pairs were formed in risky sites, but there were no differences in female sizes and progeny number, size and age near and away from scorpion burrows. Our complementary behavioral experiments revealed that bigger males stayed longer near safe burrows, and won more male-male contests than smaller conspecifics. Our findings highlight that prey can anticipate future costs of predation and use this information to assess the expected reproductive value of mates.

The Middle Eastern Cousin: Comparative Venomics of Daboia palaestinae and Daboia russelii.

Among the medically most important snakes in the world, the species belonging to the genus Daboia have been attributed to the highest number of human envenomings, deaths and disabilities. Given their significant clinical relevance, the venoms of Russell's vipers (D. russelii and D. siamensis) have been the primary focus of research. In contrast, the composition, activity, ecology and evolution of venom of its congener, the Palestine viper (D. palaestinae), have remained largely understudied. Therefore, to unravel the factors responsible for the enhanced medical relevance of D. russelii in comparison to D. palaestinae, we comparatively evaluated their venom proteomes, biochemical activities, and mortality and morbidity inflicting potentials. Furthermore, the synthesis and regulation of venom in snakes have also remained underinvestigated, and the relative contribution of each venom gland remains unclear. We address this knowledge gap by sequencing the tissue transcriptomes of both venom glands of D. palaestinae, and comparatively evaluating their contribution to the secreted venom concoction. Our findings highlight the disparity in the venom composition, function and toxicities of the two Daboia species. We also show that toxin production is not partitioned between the two venom glands of D. palaestinae.

Arthropods as the Engine of Nutrient Cycling in Arid Ecosystems.

Nutrient dynamics in most terrestrial ecosystems are regulated by moisture-dependent processes. In drylands, nutrient dynamics are often weakly associated with annual precipitation, suggesting that other factors are involved. In recent years, the majority of research on this topic focused on abiotic factors. We provide an arthropod-centric framework that aims to refocus research attention back on the fundamental role that macro-arthropods may play in regulating dryland nutrient dynamics. Macro-arthropods are prevalent in drylands and include many detritivores and burrowing taxa that remain active during long dry periods. Macro-arthropods consume and process large quantities of plant detritus and transport these nutrients to the decomposer haven within their climatically buffered and nutritionally enriched burrows. Consequently, arthropods may accelerate mineralization rates and generate a vertical nutrient recycling loop (VRL) that may assist in explaining the dryland decomposition conundrum, and how desert plants receive their nutrients when the shallow soil is dry. The burrowing activity of arthropods and the transportation of subterranean soil to the surface may alter the desert microtopography and promote desalinization, reducing resource leakage and enhancing productivity and species diversity. We conclude that these fundamental roles and the arthropods' contribution to nutrient transportation and nitrogen fixation makes them key regulators of nutrient dynamics in drylands.

Non-consumptive predator effects on prey population size: A dearth of evidence.

There is a large and growing interest in non-consumptive effects (NCEs) of predators. Diverse and extensive evidence shows that predation risk directly influences prey traits, such as behaviour, morphology and physiology, which in turn, may cause a reduction in prey fitness components (i.e. growth rate, survival and reproduction). An intuitive expectation is that NCEs that reduce prey fitness will extend to alter population growth rate and therefore population size. However, our intensive literature search yielded only 10 studies that examined how predator-induced changes in prey traits translate to changes in prey population size. Further, the scant evidence for risk-induced changes on prey population size have been generated from studies that were performed in very controlled systems (mesocosm and laboratory), which do not have the complexity and feedbacks of natural settings. Thus, although likely that predation risk alone can alter prey population size, there is little direct empirical evidence that demonstrates that it does. There are also clear reasons that risk effects on population size may be much smaller than the responses on phenotype and fitness components that are typically measured, magnifying the need to show, rather than infer, effects on population size. Herein we break down the process of how predation risk influences prey population size into a chain of events (predation risk affects prey traits, which affect prey fitness components and population growth rate, which affect prey population size), and highlight the complexity of each transition. We illustrate how the outcomes of these transitions are not straightforward, and how environmental context strongly dictates the direction and magnitude of effects. Indeed, the high variance in prey responses is reflected in the variance of results reported in the few studies that have empirically quantified risk effects on population size. It is therefore a major challenge to predict population effects given the complexity of how environmental context interacts with predation risk and prey responses. We highlight the critical need to appreciate risk effects at each level in the chain of events, and that changes at one level cannot be assumed to translate into changes in the next because of the interplay between risk, prey responses, and the environment. The gaps in knowledge we illuminate underscore the need for more evidence to substantiate the claim that predation risk effects extend to prey population size. The lacunae we identify should inspire future studies on the impact of predation risk on population-level responses in free-living animals.

Burrowing detritivores regulate nutrient cycling in a desert ecosystem.

Nutrient cycling in most terrestrial ecosystems is controlled by moisture-dependent decomposer activity. In arid ecosystems, plant litter cycling exceeds rates predicted based on precipitation amounts, suggesting that additional factors are involved. Attempts to reveal these factors have focused on abiotic degradation, soil-litter mixing and alternative moisture sources. Our aim was to explore an additional hypothesis that macro-detritivores control litter cycling in deserts. We quantified the role different organisms play in clearing plant detritus from the desert surface, using litter baskets with different mesh sizes that allow selective entry of micro-, meso- or macrofauna. We also measured soil nutrient concentrations in increasing distances from the burrows of a highly abundant macro-detritivore, the desert isopod Hemilepistus reaumuri. Macro-detritivores controlled the clearing of plant litter in our field site. The highest rates of litter removal were measured during the hot and dry summer when isopod activity peaks and microbial activity is minimal. We also found substantial enrichment of inorganic nitrogen and phosphorous near isopod burrows. We conclude that burrowing macro-detritivores are important regulators of litter cycling in this arid ecosystem, providing a plausible general mechanism that explains the unexpectedly high rates of plant litter cycling in deserts.

Observation resolution critically influences movement-based foraging indices.

Movement-based indices such as moves per minute (MPM) and proportion time moving (PTM) are common methodologies to quantify foraging behavior. Hundreds of studies have reported these indices, many without specifying the temporal resolution of their original data, and others using varying resolutions. This was done despite the likelihood that observation resolution can affect MPM and PTM estimates. Our goal was to empirically determine the sensitivity of these foraging indices to changes in the temporal resolution of the observation. We used a high-speed camera to record movement sequences of 20 Acanthodactylus boskianus lizards. Then, we gradually decreased the resolution of the data and calculated the foraging indices at different temporal resolutions. When considering the range of temporal resolutions that are relevant for field observations with unassisted vision, we found 68% and 48% difference in MPM and PTM estimates, respectively. When using the highest resolution, our estimate of MPM was an order of magnitude higher than all prior reported values for lizards. Our results raise major concerns regarding the use of already published movement-based indices, and enable us to recommend how new foraging data should be collected.

Ecological Stoichiometry: A Link Between Developmental Speed and Physiological Stress in an Omnivorous Insect.

The elemental composition of organisms belongs to a suite of functional traits that may adaptively respond to fluctuating selection pressures. Life history theory predicts that predation risk and resource limitations impose selection pressures on organisms' developmental time and are further associated with variability in energetic and behavioral traits. Individual differences in developmental speed, behaviors and physiology have been explained using the pace-of-life syndrome (POLS) hypothesis. However, how an organism's developmental speed is linked with elemental body composition, metabolism and behavior is not well understood. We compared elemental body composition, latency to resume activity and resting metabolic rate (RMR) of western stutter-trilling crickets (Gryllus integer) in three selection lines that differ in developmental speed. We found that slowly developing crickets had significantly higher body carbon, lower body nitrogen and higher carbon-to-nitrogen ratio than rapidly developing crickets. Slowly developing crickets had significantly higher RMR than rapidly developing crickets. Male crickets had higher RMR than females. Slowly developing crickets resumed activity faster in an unfamiliar relative to a familiar environment. The rapidly developing crickets did the opposite. The results highlight the tight association between life history, physiology and behavior. This study indicates that traditional methods used in POLS research should be complemented by those used in ecological stoichiometry, resulting in a synthetic approach that potentially advances the whole field of behavioral and physiological ecology.

Consequences of prey exoskeleton content for predator feeding and digestion: black widow predation on larval versus adult mealworm beetles.

Predators often feed on a wide range of prey that can vary in behavior, morphology, and physiology. The net benefits that predators gain from prey are likely related to both prey nutrient content and prey morphology or defenses. For invertebrates, the exoskeleton is a morphological trait that varies widely among species and during ontogeny and could affect nutrient extraction by predators. The goal of this study was to determine how prey exoskeleton content affected predator nutrient intake, assimilation, and excretion by comparing spiders feeding on either larval or adult mealworms of similar size. We found that the proportion of prey energy invested in digestion was greatest in spiders consuming adult mealworm beetles which had higher amounts of exoskeleton than larvae. Further, spiders extracted a greater proportion of elements, macronutrients, and energy from the larval mealworms, which had lower amounts of exoskeleton. Interestingly, total nitrogen content of prey was not a predictor of nitrogen assimilation as spiders assimilated more nitrogen from the larval mealworms, which had lower total nitrogen content. While adult beetles had higher total nitrogen content, their discarded remains of prey had large amounts of nitrogen that was nutritionally unavailable for spiders (i.e., exoskeleton). These results suggest that prey exoskeleton can affect assimilation efficiency by predators, and that a combination of macronutrient and elemental analyses may be needed to examine the quality of prey for predators and the potential consequences of predation for nutrient flows (e.g., consumer assimilation, egestion, and excretion) in ecosystems.

Considerations Used by Desert Isopods to Assess Scorpion Predation Risk.

Animals adjust behaviors to balance changes in predation risk against other vital needs. Animals must therefore collect sensory information and use a complex risk-assessment process that estimates risks and weighs costs and benefits entailed in different reactions. Studying this cognitive process is challenging, especially in nature, because it requires inferring sensory abilities and conscious decisions from behavioral reactions. Our goal was to address this empirical challenge by implementing psychophysical principles to field research that explores considerations used by desert isopods (Hemilepistus reaumuri) to assess the risk of scorpions that hunt exclusively from within their burrows. We introduced various combinations of chemical and physical cues to the vicinity of isopod burrows and recorded their detailed reactions on first encountering the cues. The isopods reacted defensively to scorpion odor but only when accompanied with excavated soil or other odors typically found near scorpion burrows. Isopods also reacted defensively to piles of excavated soil without scorpion olfactory cues, suggesting that isopods take precautions even against physical disturbances that do not necessarily reflect predator activity. Simultaneous presence of different cues provoked graded responses, possibly reflecting an additive increase in risk estimation. We conclude that wild isopods use defensive reactions toward environmental signals only when the integrated perceptual information implies an active scorpion burrow or when they lack data to refute this possibility.

How to use (and not to use) movement-based indices for quantifying foraging behaviour.

Movement-based indices such as moves per minute (MPM) and proportion time moving (PTM) are common methodologies to quantify foraging behaviour. We explore fundamental drawbacks of these indices that question the ways scientists have been using them and propose new solutions.To do so, we combined analytical and simulation models with lizards foraging data at the individual and species levels.We found that the maximal value of MPM is constrained by the minimal durations of moves and stops. As a result, foragers that rarely move and those that rarely stop are bounded to similar low MPM values. This implies that (1) MPM has very little meaning when used alone, (2) MPM and PTM are interdependent, and (3) certain areas in the MPM-PTM plane cannot be occupied. We also found that MPM suffers from inaccuracy and imprecision.We introduced a new bias correction formula for already published MPM data, and a novel index of changes per minute (CPM) that uses the frequency of changes between move and stop bouts. CPM is very similar to MPM, but does not suffer from bias. Finally, we suggested a new foraging plane of average move and average stop durations. We hope that our guidelines of how to use (and not to use) movement-based indices will add rigor to the study of animals' foraging behaviour.

Trophic cascade alters ecosystem carbon exchange.

Trophic cascades--the indirect effects of carnivores on plants mediated by herbivores--are common across ecosystems, but their influence on biogeochemical cycles, particularly the terrestrial carbon cycle, are largely unexplored. Here, using a (13)C pulse-chase experiment, we demonstrate how trophic structure influences ecosystem carbon dynamics in a meadow system. By manipulating the presence of herbivores and predators, we show that even without an initial change in total plant or herbivore biomass, the cascading effects of predators in this system begin to affect carbon cycling through enhanced carbon fixation by plants. Prolonged cascading effects on plant biomass lead to slowing of carbon loss via ecosystem respiration and reallocation of carbon among plant aboveground and belowground tissues. Consequently, up to 1.4-fold more carbon is retained in plant biomass when carnivores are present compared with when they are absent, owing primarily to greater carbon storage in grass and belowground plant biomass driven largely by predator nonconsumptive (fear) effects on herbivores. Our data highlight the influence that the mere presence of predators, as opposed to direct consumption of herbivores, can have on carbon uptake, allocation, and retention in terrestrial ecosystems.

Linking predation risk, herbivore physiological stress and microbial decomposition of plant litter.

The quantity and quality of detritus entering the soil determines the rate of decomposition by microbial communities as well as recycle rates of nitrogen (N) and carbon (C) sequestration. Plant litter comprises the majority of detritus, and so it is assumed that decomposition is only marginally influenced by biomass inputs from animals such as herbivores and carnivores. However, carnivores may influence microbial decomposition of plant litter via a chain of interactions in which predation risk alters the physiology of their herbivore prey that in turn alters soil microbial functioning when the herbivore carcasses are decomposed. A physiological stress response by herbivores to the risk of predation can change the C:N elemental composition of herbivore biomass because stress from predation risk increases herbivore basal energy demands that in nutrient-limited systems forces herbivores to shift their consumption from N-rich resources to support growth and reproduction to C-rich carbohydrate resources to support heightened metabolism. Herbivores have limited ability to store excess nutrients, so stressed herbivores excrete N as they increase carbohydrate-C consumption. Ultimately, prey stressed by predation risk increase their body C:N ratio, making them poorer quality resources for the soil microbial pool likely due to lower availability of labile N for microbial enzyme production. Thus, decomposition of carcasses of stressed herbivores has a priming effect on the functioning of microbial communities that decreases subsequent ability to of microbes to decompose plant litter. We present the methodology to evaluate linkages between predation risk and litter decomposition by soil microbes. We describe how to: induce stress in herbivores from predation risk; measure those stress responses, and measure the consequences on microbial decomposition. We use insights from a model grassland ecosystem comprising the hunting spider predator (Pisuarina mira), a dominant grasshopper herbivore (Melanoplus femurrubrum),and a variety of grass and forb plants.

Predation risk, stoichiometric plasticity and ecosystem elemental cycling.

It is widely held that herbivore growth and production is limited by dietary nitrogen (N) that in turn constrains ecosystem elemental cycling. Yet, emerging evidence suggests that this conception of limitation may be incomplete, because chronic predation risk heightens herbivore metabolic rate and shifts demand from N-rich proteins to soluble carbohydrate-carbon (C). Because soluble C can be limiting, predation risk may cause ecosystem elemental cycling rates and stoichiometric balance to depend on herbivore physiological plasticity. We report on a stoichiometrically explicit ecosystem model that investigates this problem. The model tracks N, and soluble and recalcitrant C through ecosystem compartments. We evaluate how soluble plant C influences C and N stocks and flows in the presence and absence of predation risk. Without risk, herbivores are limited by N and respire excess C so that plant-soluble C has small effects only on elemental stocks and flows. With predation risk, herbivores are limited by soluble C and release excess N, so plant-soluble C critically influences ecosystem elemental stocks flows. Our results emphasize that expressing ecosystem stoichiometric balance using customary C:N ratios that do not distinguish between soluble and recalcitrant C may not adequately describe limitations on elemental cycling.