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Biocrusts are key ecosystem engineers that are being destroyed due to anthropogenic disturbances such as trampling, agriculture and mining. In hyper-arid regions of the Negev Desert, phosphate has been mined for over six decades, altering the natural landscape over large spatial scales. In recent years, restoration-oriented practices were mandated in mining sites, however, the impact of such practices on the ecosystem, particularly the biocrust layer, has not been tested. Here, we evaluated post-mining biocrust bacterial communities and compared them to undisturbed (reference) biocrusts. We collected samples from four mining sites (each restored at a different year) and their corresponding reference sites. We hypothesized that post-mining bacterial communities would differ significantly from reference communities, given the slow regeneration of the biocrust. We also hypothesized that bacterial communities would vary among post-mining plots based on their restoration age. To test these hypotheses, we assessed the abundance and diversity of bacterial communities by sequencing the 16S rDNA and their photosynthetic potential by quantifying the abundance of cyanobacteria and chlorophyll a. The bacterial diversity was lower, and community composition differed significantly between post-mining and reference biocrusts. In addition, cyanobacteria abundances and chlorophyll a content were lower in post-mining biocrusts, indicating lower photosynthetic potential. However, no significant changes in bacterial communities were detected, regardless of the restoration age. We suggest that the practices implemented in the Negev mines may not support the recovery of the biocrust bacterial communities, particularly the cyanobacteria. Thus, active restoration measures are needed to accelerate the regeneration time of biocrusts at the hyper-arid Negev mines.

To understand patterns of alpha, beta and gamma diversities in fragmented landscapes we need to explore the three scale components in relation to potential drivers in a scale-dependent manner. Often, the drivers themselves can be partitioned to alpha, beta and gamma diversities. Thus, one can hypothesize that the scale-components of species diversity and drivers’ diversity match, i.e., that species alpha diversity is mainly explained by drivers’ alpha diversity, beta by beta and gamma by gamma. Here, we explore this ‘scale-matching’ hypothesis for spiders in two fragmented agricultural landscapes. In each landscape, we sampled spiders and their potential prey in 12 patches. Then, we sub-sampled pseudo-landscapes in which we calculated spider alpha, beta and gamma diversities using multiplicative diversity-partitioning. Next, we used variance partitioning analysis to explore the relative contribution of eleven explanatory variables from five thematic groups (sampling intensity, area, connectivity, habitat diversity and prey diversity), while further partitioning the habitat and prey diversities to their corresponding alpha, beta and gamma diversities. We found considerable evidence for scale-matching, with spiders’ alpha and beta diversities explained mostly by the corresponding alpha and beta diversities (respectively) of prey and/or habitat. We further found a strong effect of connectivity on spider beta diversity, but not on alpha and gamma diversities. For spiders gamma diversity, a cross-scale effect was observed. Our results suggest that multiple drivers from multiple scales interact in structuring patterns of spider alpha, beta and gamma diversities in agro-ecosystems, yet the strongest effects are of those drivers that match in scale.

Adult males of many migratory species arrive at breeding grounds before females and young. In a 34-year study of the masked shrike, Lanius nubicus, an age- and sex-specific pattern of spring arrival was distinguished. Adult males arrived first followed by juvenile males and adult females, whereas juvenile females appeared last. We hypothesized that these differences in migratory strategies would be reflected in a differential response to climate conditions at the wintering grounds. Testing correlations between spring arrival time and winter climate conditions provided strong support to our hypothesis. Adult males’ arrival time exhibited high associations with climate conditions in early spring, upon migratory take-off, whereas juvenile males responded mostly to conditions in November, upon autumn arrival in Africa. Adult females responded to both parameters in autumn and early spring, whereas young females’ arrival correlated only with a few variables in autumn. GLM models of median spring arrival day for all categories but the young females were highly statistically significant with adjusted R-squared values of 0.81–0.93. The emerging pattern of different associations between timing of spring migration and climate conditions at the wintering grounds sheds new light on existing evolutionary theories regarding age- and sex-specific migratory strategies.

Landscape heterogeneity is a general feature of natural environments, strongly affected by habitat fragmentation. It can affect a population's dynamics and probability of extinction. Fragmentation increases among-patch isolation and decreases patch size, resulting in a reduction in available resources in smaller patches. To persist, animals must be able to translate the variation imposed by fragmentation into adaptive energy allocation strategies that enable populations to avoid extinction. This means that physiological adaptations are expected to reflect changes in landscape configuration, especially in the size of the natural habitat patches and degree of isolation among them. We propose a novel, integrative conceptual framework in which spatial characteristics of the environment, imposed by fragmentation, lead to specific life-history traits that increase survival (at the individual level) and decrease the likelihood of extinction (as an emergent property at the population level). We predict that a resource allocation trade-off between the life-history traits of reproduction and dispersal along a fragmentation gradient will emerge. Populations occurring in patches of different sizes and isolations along gradients of fragmentation and productivity will exhibit differences in the strength of the dispersal-reproduction trade-off. Emerging from this framework are several explicit and testable hypotheses that predict that the dispersal-reproduction trade-off will be shaped by landscape heterogeneity imposed by fragmentation. Hence, this trade-off serves as the mechanistic link that translates environmental variation created by fragmentation into variation in species abundances and population dynamics by lowering local extinction probability and increasing overall population persistence.

Land sharing strategies in agricultural landscapes need to allow for organisms to move between natural areas and different crops within an agro-ecological landscape in order to reduce extinction probability and the negative effects of small isolated populations. In this study, we tested whether legume or wheat fields differed in their effects on reptiles’ movement patterns. We conducted our study in an agro-ecosystem consisting of small isolated natural habitat patches nested within agricultural fields. We trapped reptiles in sampling arrays before and after harvest in both wheat and legume fields, and in adjacent natural habitat patches. For both crops, prior to harvest, we found an increase in movements of Trachylepis vittata, the most common reptile in our study, from the natural habitat patches into fields, but negligible movement in the opposite direction. In both crops before harvest, the individuals that moved into the fields were adults of better body condition than those remaining in the natural habitat patch, suggesting that long-distance movements were only possible for individuals with high prospective fitness. After harvest, no movements were documented between wheat fields and natural habitat patches. However, in legume fields, a high symmetrical movement (i.e. in both directions) of individuals of similar body condition between fields and natural habitat patches took place. Importantly, newborn lizards were only found in the natural habitat patches and in post-harvest legume fields. Our results suggest that agricultural heterogeneity, through a mixture of crop types may mitigate some of the negative effects of particular crops on biodiversity. As crop rotation between wheat and legume fields is common worldwide, our findings highlight the importance of creating an agricultural mosaic to enhance biodiversity permeability within the agricultural matrix.

This is the first population genomic analysis that examines micro-evolutionary changes in co-occurring plant species, in fragmented habitats, along a precipitation gradient. We applied Genotype-by-Sequencing (GBS) on three different annual Asteraceae on the same sites that share similar life history strategies, but different pollination and dispersal strategies (Catananche lutea, Geropogon hybridus, Urospermum picroides). We tested if the genetic diversity for all species correlates with effective population sizes along the precipitation gradient and whether therefore genetic drift drives evolution at range edges. However, our results support this hypothesis for only one species (C. lutea) but refute it for the other two. Potential genomic signals of adaptation to the high precipitation gradient were found only in one species (U. picroides). We also tested if species with complex pollination and diversified dispersal strategies are less affected by small scale habitat fragmentation, but the opposite is true. Biological differences between the species studied (pollination and dispersal) explain population genetic differences along the precipitation gradient at range edges rather than environmental filters. Pairwise correlation analyses showed no or only weak similarities in the accumulation of mutations in the different species. Our results suggest that the processes driving evolution in different co-occurring species are different, so that most genomic similarities in our investigations may only represent temporary stages in evolution.

AimPatterns that relate species richness with fragment area (the species–area relationship, SAR) and with isolation (the species–isolation relationship, SIR) are well documented. However, those that relate species density – the number of species within a standardized area – with fragment area (D-SAR) or isolation (D-SIR) have not been sufficiently explored, despite the potential for such an analysis to disentangle the underlying mechanisms of SARs and SIRs. Previous spatial theory predicts that a significant D-SAR or D-SIR is unlikely to emerge in taxa with high dispersal limitation, such as plants. Furthermore, a recent model predicts that the detection and the significance of D-SARs or D-SIRs may decrease with grain size. We combined a literature review with grain size-dependent sampling in a fragmented landscape to evaluate the prevalence and grain size-dependent nature of D-SARs and D-SIRs in plants.LocationWorldwide (review) and a semi-arid agro-ecosystem in Israel (case study).MethodsWe combined an extensive literature review of 31 D-SAR studies of plants in fragmented landscapes with an empirical study in which we analysed grain size-dependent D-SARs and D-SIRs using a grain size-dependent hierarchical sampling of species density and species richness in a fragmented, semi-arid agro-ecosystem.ResultsWe found that significantly increasing D-SARs are rare in plant studies. Furthermore, we found that the detection of a significant D-SAR is often possible only after the data have been stratified by species, habitat or landscape characteristics. The results from our case study indicated that the significance and the slopes of both D-SARs and D-SIRs increase as grain size decreases.Main conclusionsThese results call for a careful consideration of scale while analysing and interpreting the responses of species richness and species density to fragmentation. Our results suggest that grain size-dependent analyses of D-SARs and D-SIRs may help to disentangle the mechanisms that generate SARs and SIRs and may enable early detection of the effects of fragmentation on plant biodiversity.

Most papers on the physical condition of birds during spring migration focused on food availability preceding migratory take-off. Only a few studies examined the effect of climate conditions at the wintering grounds upon autumn arrival on bird physical condition later on. Here, we hypothesized that environmental conditions upon arrival at the wintering grounds, and not necessarily upon departure, have a crucial carry-over effect on bird spring migration. Using 29,000 observations of the lesser whitethroat, Sylvia curruca, and the eastern Bonelli's warbler, Phylloscopus orientalis, we found temperatures upon arrival at the African wintering grounds to be the only climatic variable correlated with birds' body state upon spring stopover in Israel, six months later. Two different mechanisms could explain these results. One possibility is that high temperatures create favorable conditions for insect activity, which allows rapid recovery from autumn migration and hence successful winter survival and maintenance. Another possible scenario is that harsh conditions, due to the heat and dry environment, cause high mortality, permitting survival of larger individuals which, then, enjoy reduced inter- and intra-specific competition. Whatever the mechanism is, our findings suggest that conditions upon autumn arrival, and not necessarily at the end of winter as traditionally thought, may have a major impact on migrating birds.

Intensive agricultural activity over large areas on earth, which is necessary to meet the increasing demand of a growing human population, may lead to biodiversity loss. This loss may be mitigated by keeping natural and semi-natural patches within agricultural fields to allow the maintenance of biological diversity (‘Wildlife Friendly Agriculture’). We conducted our study in an agroecosystem comprised of small isolated patches nested within agricultural fields. We trapped reptiles in 13 sampling sites, each of which included arrays of pitfall traps in a natural patch, in the adjacent wheat field and at the patch-field edge. We conducted six trapping sessions in the spring – four times before, once immediately after and once a week after the wheat harvest. Prior to the harvest, we found an intensive movement of Trachylepis vittata, the most common reptile in our study, from the semi-natural patches into the fields, but negligible movement in the opposite direction. This pre-harvest directional movement corresponded with higher abundance of prey (i.e., arthropods) in the wheat field compared to the natural patches in early spring. The individuals that moved into the fields were adults of better body condition than those remaining in the patch, suggesting that the motivation for movement was habitat preference by individuals with high prospective fitness rather than the exclusion of subordinates. The population of T. vittata in the wheat fields and movement across habitats dropped to zero during and after the harvest. Our results provide strong evidence that the agricultural fields serve as an ecological trap to organisms inhabiting nearby natural habitats. We suggest that plans for Wildlife-Friendly Agriculture for biodiversity conservation should consider also potential negative effects, such as the ecological trap effect.

Spatial environmental heterogeneity is considered a fundamental factor for the maintenance of plant species richness. However, it still remains unclear whether heterogeneity may also facilitate coexistence at fine grain sizes or whether other processes, like mass effects and source sink dynamics due to dispersal, control species composition and diversity at these scales. In this study, we used two complimentary analyses to identify the role of heterogeneity within 15m×15m plots for the coexistence of species-rich annual communities in a semi-arid environment along a steep precipitation gradient. Specifically, we: (a) analyzed the effect of environmental heterogeneity on species, functional and phylogenetic diversity within microsites (alpha diversity, 0.06m2 and 1m2), across microsites (beta diversity), and diversity at the entire plot (gamma diversity); (b) further we used two null models to detect non-random trait and phylogenetic patterns in order to infer assembly processes, i.e. whether co-occurring species tend to share similar traits (trait convergence) or dissimilar traits (trait divergence). In general, our results showed that heterogeneity had a positive effect on community diversity. Specifically, for alpha diversity, the effect was significant for functional diversity, and not significant for either species or phylogenetic diversities. For beta diversity, all three measures of community diversity (species, functional, and phylogenetic) increased significantly, as they also did for gamma diversity, where functional measures were again stronger than for species or phylogenetic measures. In addition, the null model approach consistently detected trait convergence, indicating that species with similar traits tended to co-occur and had high abundances in a given microsite. While null model analysis across the phylogeny partly supported these trait findings, showing phylogenetic underdispersion at the 1m2 grain size, surprisingly when species abundances in microsites were analyzed they were more evenly distributed across the phylogenetic tress than expected (phylogenetic overdispersion). In conclusion, our results provide compelling support that environmental heterogeneity at a relatively fine scale is an important factor for species co-existence as it positively affects diversity as well as influences species assembly. Our study underlines the need for trait-based approaches conducted at fine grain sizes in order to better understand species coexistence and community assembly. Spatial environmental heterogeneity is considered a fundamental factor for the maintenance of plant species richness. However, it still remains unclear whether heterogeneity may also facilitate coexistence at fine grain sizes or whether other processes, like mass effects and source sink dynamics due to dispersal, control species composition and diversity at these scales. In this study, we used two complimentary analyses to identify the role of heterogeneity within 15 m × 15 m plots for the coexistence of species-rich annual communities in a semi-arid environment along a steep precipitation gradient. Specifically, we: (a) analyzed the effect of environmental heterogeneity on species, functional and phylogenetic diversity within microsites (alpha diversity, 0.06 m2 and 1 m2), across microsites (beta diversity), and diversity at the entire plot (gamma diversity); (b) further we used two null models to detect non-random trait and phylogenetic patterns in order to infer assembly processes, i.e. whether co-occurring species tend to share similar traits (trait convergence) or dissimilar traits (trait divergence). In general, our results showed that heterogeneity had a positive effect on community diversity. Specifically, for alpha diversity, the effect was significant for functional diversity, and not significant for either species or phylogenetic diversities. For beta diversity, all three measures of community diversity (species, functional, and phylogenetic) increased significantly, as they also did for gamma diversity, where functional measures were again stronger than for species or phylogenetic measures. In addition, the null model approach consistently detected trait convergence, indicating that species with similar traits tended to co-occur and had high abundances in a given microsite. While null model analysis across the phylogeny partly supported these trait findings, showing phylogenetic underdispersion at the 1m2 grain size, surprisingly when species abundances in microsites were analyzed they were more evenly distributed across the phylogenetic tress than expected (phylogenetic overdispersion). In conclusion, our results provide compelling support that environmental heterogeneity at a relatively fine scale is an important factor for species co-existence as it positively affects diversity as well as influences species assembly. Our study underlines the need for trait-based approaches conducted at fine grain sizes in order to better understand species coexistence and community assembly.

Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. −30% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N- and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height towards aridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (≫50 is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors. Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. −30% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N- and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height towards aridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (≫50%) is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors.

Numerous studies report shifts in bird migration phenology, presumably owing to global warming. However, most studies focus on migration patterns in the Northern Hemisphere. In this study, we investigated associations between weather conditions in African wintering grounds of the lesser whitethroat, Sylvia curruca, and spring arrival time in Eilat, Israel. Using multivariate regression models, we analysed a 30-year dataset in order to examine correlations between median springtime arrival and 46 climate variables of the wintering quarters. The model obtained exhibited a highly statistical fit, involving mean precipitation in February and March with negative effects and number of wet days during November–February. February precipitation levels were also the major factor associated with the interquartile range of arrival time. Interestingly and contrary to published results, annual or seasonal precipitation showed no correlation with spring arrival time, nor did temperature. Moreover, winter in this region falls into dry season with negligible rainfall quantities. Hence, it is unlikely that precipitation effect on habitat productivity is a driving force of migration, as suggested by other studies. Instead, we propose that precipitation in February acts as a cue for the birds, indicating the approach of spring and migration time.

Dominance relationships among species play a major role in the structure of animal communities. Yet, dominant species with different trade-offs in resource exploitation and monopolization could affect community structure in variable ways. In ants, dominant species could be classified into either behavioral dominants that exhibit territorial aggression or numerical dominants that exhibit high biomass or frequency of occurrence. While each class of dominance has generally been found to negatively affect the foraging activity of species in ant communities, the concurrent effect of both classes of species has never been tested. Here, we examined the effects of two behaviorally dominant species, Crematogaster inermis and Monomorium salomonis, and a numerically dominant species, Messor arenarius, on the foraging behavior of seed-eating species in a desert ant assemblage. In a 1-year study, the foraging activity of the ant species was assessed using seed baits, which were sampled during night and day. While the numerically dominant species exhibited high foraging efficiency and negatively affected the ability of other seed-eating species to obtain seeds, significantly more seeds remained at baits that were occupied the previous night by each of the two behaviorally dominant species, possibly due to aggressive exclusion of M. arenarius foragers from the baits. This exclusion also facilitated greater foraging activity of the seed-eating species. Our results demonstrate how these two types of dominance could differently affect the foraging activity of ant species in the community.

Aims: (1) Understanding how the relationship between species richness and its determinants depends on the interaction between scales at which the response and explanatory variables are measured. (2) Quantifying the relative contributions of local, intermediate and large-scale determinants of species richness in a fragmented agro-ecosystem. (3) Testing the hypothesis that the relative contribution of these determinants varies with the grain size at which species richness is measured. Location: A fragmented agro-ecosystem in the Southern Judea Lowland, Israel, within a desert–Mediterranean transition zone. Methods: Plant species richness was estimated using hierarchical nested sampling in 81 plots, positioned in 38 natural vegetation patches within an agricultural matrix (mainly wheat fields) among three land units along a sharp precipitation gradient. Explanatory variables included position along that gradient, patch area, patch isolation, habitat heterogeneity and overall plant density. We used general linear models and hierarchical partitioning of variance to test and quantify the effect of each explanatory variable on species richness at four grain sizes (0.0625, 1, 25 and 225 m2). Results: Species richness was mainly affected by position along a precipitation gradient and overall plant density, and to a lesser extent by habitat heterogeneity. It was also significantly affected by patch area and patch isolation, but only for small grain sizes. The contribution of each explanatory variable to explained variance in species richness varied with grain size, i.e. scale-dependent. The influence of geographic position and habitat heterogeneity on species richness increased with grain size, while the influence of plant density decreased with grain size. Main conclusions: Species richness is determined by the combined effect of several scale-dependent determinants. Ability to detect an effect and effect size of each determinant varies with the scale (grain size) at which it is measured. The combination of a multi-factorial approach and multi-scale sampling reveals that conclusions drawn from studies that ignore these dimensions are restricted and potentially misleading.

To test the effects of connectivity on fish diversity in isolated reef patches we deployed pairs of artificial reefs (AR) at constant distances (12 and 25 m) from a large continuous reef and added series of small ARs to half of them. These small reefs served as stepping-stones to increase fish movement between the large ARs and the continuous reef. Species gain and loss curves were compared to obtain deeper understanding of the underlying mechanisms in relation to the theory of island biogeography. We found that AR without stepping-stones maintained up to 57-fold more individuals than ARs with stepping-stones during mass recruitment events that were dominated by the family Apogonidae. However, in-between these mass recruitment events the overall impact of the stepping-stones on richness and abundance was small. By contrast, increasing distance from the continuous reef from 12 to 25 m increased fish richness, although this was confounded with time since AR deployment. Increase in richness with distance was predominantly caused by elevated species gain. Our results suggest that fish assemblages respond in distinct ways to two types of isolation: (1) Increased richness with distance from a continuous reef, and (2) increased fluctuation in abundance with decreased connectivity. The latter may be the outcome of accumulation and then abrupt emigrations of fishes from isolated reefs. This study confirms that the spatial setting of reef patches in relation to larger or continuous reefs alters fish assemblages, which may be important for planning AR deployments that maximizes diversity.

Fish abundance is often better predicted by microhabitat variables on continuous reefs than on isolated patch reefs. Although this was suggested to stem from reduced post-recruitment relocation, this has not been shown experimentally. We found the relationship between the presence of a coral-dwelling fish, Dascyllus marginatus, and the size of its coral host to differ between corals on continuous reefs and the sparsely distributed corals on sandy bottoms. Empty transplanted corals were colonized exclusively by new recruits when on the sandy bottom, and both by new recruits and post-recruitment dispersal of adults when on the continuous reef. New recruits settled predominantly into small corals, although analyses of recruitment patterns were confounded by low recruitment in the studied years. Both tank experiments and field survey data suggest that the presence of recruits in small corals is at least partially driven by predation by the dottyback, Pseudochromis olivaceus, which lives predominantly in large corals within both habitats. Consequently, we suggest that the relationship between fish presence and coral size differs between the habitats due to coral size dependent predation on recruits and variability in the importance of direct recruitment to replenish fish populations.

Many agricultural greenhouse crops suffer reduced yields due to insufficient pollination. This problem can be alleviated by introducing efficient pollinating insects into the greenhouse. The bee Xylocopa puhescens Spinola 1838, a candidate for domestication as an agricultural pollinator, is unique in its facultative social organization. Females either nest solitarily, or together with a second female (a non-reproducing guard). Social nesting occurs when food and nest sites are limited, and carries fitness benefits and costs to the bees as.compared to solitary nesting. The implications of X pubescens' social organization for crop pollination were investigated. Honeydew melons were grown as a model crop in a small greenhouse. The non-crop plants Portulaca oleracea L, Solanum rantonnetii C. Lavandula angustifolia Mill and Ocimum basilicum L supplemented the bees' diet. Social and solitary X pubescens nesters were introduced into the greenhouse in alternation. The bees' daily activity pattern, the frequency and duration of visits to each flower species, and the run-lengths of consecutive visits to each flower species were recorded. The melons' fruit set, and the fruits' mass and seed number, were determined. Social nesters visited P oleracea more frequently than solitary bees when this species was in bloom. After P oleracea finished blooming, socially nesting bees visited melon more often than solitary nesters Social bees spent a longer time at the melon patch and tended to be more flower constant than solitary nesters, but spent less time per flower than solitary individuals. Solitary and social bees did not differ in their daily activity patterns and flower visitation rates. Pollination by both types of nesters resulted in similar fruit sets, fruit mass and fruit seed numbers. The dissimilarities in foraging behaviour may reflect differences in the dietary demands of solitary vs social nesters. The similarity in fruit sets and flower constancy suggests that both nest types provide pollination services of similar quality.

The peace treaty between Israel and Jordan found the Arava desert ecosystem, shared by the two countries, in a state of developmental dichotomy. On the Israeli side, vast lands have been settled and transformed into agricultural fields, while the Jordanian side has remained relatively intact and inhabited by only a few traditional and pastoral societies. This study examined the effect of different landscape units and proximity to agriculture on reptile diversity and rodent community structure on both sides of the border. It appears that in addition to the effect of proximity to agricultural fields and landscape habitat, the border between the two countries may play a role in determining diversity on the respective sides. While reptile abundance was generally higher on the Israeli side of the border, diversity was found to be significantly higher on the Jordanian side. Rodent community structure also revealed significant differences between the two sides of the border, mainly due to the more favorable conditions for psammophilic gerbils in Jordan. When comparing Western society with pastoral traditional society, it appears that development activities of the former have altered diversity and community structure of the taxa studied in the Arava. We suggest that in addition to the effects of habitats and human disturbances, such as modern agricultural practices, cultural differences between societies should be considered when conservation plans are developed for cross-border ecosystems.

Summary 1. We tested the hypothesis that Gerbillus henleyi (de Winton 1903), the smallest species (10 g) of a pssamophilic guild in Israel, is scarce on relatively productive dunes of the Israeli desert, due to negative interactions from the common G. allenby i (Thomas 1918) and G. pyramidum (Geoffroy 1825). 2. The alternative hypothesis was that scarcity on sand resulted from the size of its naked hind feet, that do not allow efficient locomotion on sand. 3. Despite their naked soles the weight-bearing surface of G. henleyi feet carry less mass/area than those of any other species. 4. We measured interaction coefficients with the two common species using fieldmanipulation experiments in two enclosures. 5. Habitat usage of G. henleyi changed from significantly preferring the stabilized sand, when alone, to significantly using the semistabilized dune, when G. allenbyi was also present. 6. We also estimated the interaction coefficients and calculated the G. henleyi ’s isoclines competing with the two common gerbil species using a technique we developed elsewhere. 7. The stability analysis of the isoclines of G. henleyi competing with either G. allenbyi or with G. pyramidum suggests that stable coexistence occurs when G. henleyi is relatively scarce while the competitors are common. 8. Interspecific competition from either G. allenbyi or G. pyramidum accounts for 90·3% reduction in G. henleyi density, relative to when it is alone. 9. We concluded that the negative interactions from congeners was the major cause for the scarcity of G. henleyi on the relatively rich sand dunes of the Israeli desert.

We studied the large-scale spatialÐtemporal distribution pattern of the Mediterranean fruit sy, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) to understand whether long-term dispersal may affect its maintenance in a highly patchy and isolated arid environment. The study area contained few dispersed settlements and agricultural plots covering 3,000 km 2 , ranging from 100 to 900 m in altitude in southern Israel. Due to the arid nature of the study area, no wild hosts existed outside the settlements. Data were collected from three settlements at different altitudes from 1999 to 2002. During 2000, adult sies were monitored throughout the entire area. Data were incorporated into a geographic information system, and weekly spatial distribution charts were produced and analyzed. Our results suggest that, within the study area, C. capitata overwinters in sites 400 mi n altitude with abundant winter hosts but not at higher altitudes. In remote settlements and agricultural areas at higher altitudes, sies reappeared every year in midsummer, which can only be explained by annual migration from the lower altitudes. Our results demonstrate that sies migrate at least 50 km, probably over a single generation, before reaching some of the remote havens within the study area. This long-distance migration has far-reaching consequences for maintenance of remote populations and should be taken into consideration for future basic research and applied control practices of C. capitata.

A positive correlation is often found between fish species diversity on coral reefs and their degree of isolation. We examined whether isolation creates differences in fish assemblages among reefs through changes in predation pressure. First, small artificial reefs were placed at increasing distances from a naturally continuous reef, over a sloping bottom. Species richness and density of each species increased with isolation. Next, artificial reefs were relocated, together with all their fish inhabitants, closer to the natural reef. Following relocation, resident fishes of the artificial reefs exhibited a sharp decline in numbers, while predatory fish density increased. Both video obser- vations of the relocated artificial reefs and the correlation between piscivore aggregation and decline of the resident fishes revealed that the cause of decline was the strong predation around the artificial reefs. The decline was density-dependent, such that the per-capita rate of decline was higher on arti- ficial reefs with higher fish density. This study shows that the impact of piscivores on resident fish is modified by small-scale isolation. The results suggest that the high density of fish on small, isolated reefs is enabled by low predation.

The geographical distribution of Mediterranean fruit fly, Ceratitis capitata (Wiedemann), is limited to relatively warm regions and does not cross the 41° north latitude. A recent study has demonstrated that, in contrast to the earlier suggested hypothesis, that Mediterranean fruit fly can survive the winter in relatively cold areas, it does not overwinter in such areas but reinvades them annually from adjacent warmer sites. In the present work, we use a large-scale spatial approach to describe the spatiotemporal dynamics of Mediterranean fruit fly in a large heterogeneous landscape as well as discuss the consequences of such dynamics for basic research and control practices. During a 2-yr period (2000–2001), over the central part of Israel (6,875 km2), adult males were monitored and recorded weekly by using Steiner-like traps. Data obtained were incorporated into a geographic information system to produce weekly distribution maps of C. capitata within the study area. The patterns of change in population sizes through time were consistent in both years, suggesting that C. capitata overwinters along the coastal plain and the Jordan Valley. Reinvasion to the higher altitudes is followed by a population increase in the warmer, lower altitudes, during the spring and early summer. By midsummer, flies were found at all sites within the study area. However, flies started disappearing from the high-altitude areas during fall and were absent from these areas during winter. We suggest that understanding such spatiotemporal dynamics has far-reaching consequences for future basic research as well as control and eradication programs.

Contrary to the past belief that the species richness-productivity relationship (SRPR) is characterized by a single “true” pattern, several analyses have shown that a heterogeneous distribution of SRPRs exists and that the patterns' distribution of a SRPR is sensitive to the directness of the estimates of productivity. A simple conceptual model demonstrates that taking into account underlying relationships connecting productivity with species richness produces a heterogeneous distribution of relationships. We suggest that the search for a single pattern for SRPR has reached a dead end, and that we should direct our research focus to exploring the mechanisms responsible for the various SRPRs.

We studied the effect of a dominant species, Gerbillus pyramidum (Egyptian sand gerbil), on the patch use of its subordinate competitor, G. andersoni allenbyi (Allenby's gerbil), to better understand interspecific competition between the two species. We used manipulated resource patches (seed trays) covered with cages with two adjustable species-specific gates (either opened or closed to the bigger-dominant species, but always opened to the subordinate one). We recorded species tracks around and on the seed trays and giving-up densities (GUDs) of seeds in the trays after each night of foraging. G. a. allenbyi depleted seed patches to a lower level whenever G. pyramidum was given the opportunity to forage on the seed trays (i.e., present on the grid). This result held regardless of whether G. pyramidum was actually present at a particular station. We suggest that competition from G. pyramidum occurs both directly by interference, in which G. a. allenbyi is forced to be active in the late part of the night, and indirectly by exploitation via resource depletion by G. pyramidum in the early part of the night. The results suggest that interspecific competition from G. pyramidum reduces seed availability and the richness of the environment for G. a. allenbyi enough to affect the marginal value of energy for G. a. allenbyi individuals and cause them to experience lower costs of predation and manifest lower GUDs.

Large-scale ecological patterns emerge from multiple biological processes that interact on several scales and are affected by the heterogeneity of the environment. I use a landscape simulation model to explore and analyze how interactions among various ecological processes affect patterns of mammalian species diversity according to body size. The model incorporates ecological realism by using mammalian allometric relationships with realistic values taken from field data. It uses energy as a common currency to model and compare how species of different sizes use their environment. Components of the model include landscape, habitat, patch, community, species, and population. Processes include continuous-time population growth, saturation effect, species-habitat matching, demographic stochasticity (local-scale), dispersal, and catastrophic stochasticity (landscape-scale). I simulated a 16-patch landscape, with each patch having a unique habitat, and a 10-species pool in which species differ only in body size. Summed over the landscape, a log-normal relationship between body size and species abundance emerges. Additionally, geographic range increases nonlinearly with the log of body size. These patterns are qualitatively consistent with patterns of mammalian species diversity observed in natural systems, suggesting that they can emerge from scaling of habitat use with body size. I also simulated a simple 4-patch landscape, with each patch having a unique habitat, and a 26-species pool to draw from in which species differ only in body size. The results show that interspecific competition by itself reduces species diversity in each habitat and in the entire landscape. Stochasticity depresses mean population sizes but opens opportunities for species to avoid competitive exclusion. It also allows different habitats to have different communities that are characterized by the consequences of which large species become locally extinct at random. Demographic and catastrophic stochasticities differ, however, in their characteristic effects on species diversity. These characteristic effects can be used as fingerprints to identify processes acting in nature. Dispersing individuals move between habitats and reestablish the local populations of their species. Overall, the model provides us with predictions of how process-interactions in heterogeneous landscape may affect species composition and community structure.

We examined the foraging strategy and quantified the foraging traits of two nocturnal rodent species, Allenby’s gerbil (Gerbillus allenbyi) and the greater Egyptian sand gerbil (Gerbillus pyramidum). In the laboratory, both species used two distinct foraging strategies: either they immediately consumed seeds found in a patch (seed tray); or they collected and delivered the seeds to their nest box for later consumption. Moreover, we found a transition in foraging strategy among individual G. allenbyi under laboratory conditions; they all began by consuming the seeds on the tray and, after 7 days on average, switched to the collecting strategy. By contrast, in the field both species used only one foraging strategy; they collected and delivered the seeds to their burrow or to surface caches for later consumption. Furthermore, G. allenbyi and G. pyramidum collected seeds at significantly higher rates in the field than in the laboratory because the seed encounter rates for both species were higher in the field. This suggests that in natural conditions, probably involving predation risk and competitive pressure, gerbils must respond in two ways: (1) they must choose a foraging strategy that reduces predation risk by minimizing time spent feeding outside their burrows; and (2) they must forage more efficiently. In the field, seed handling time of the larger species, G. pyramidum, was shorter than that of the smaller one, G. allenbyi. This difference may give G. pyramidum an advantage when resource levels are high and when most of a forager’s time is spent handling seeds rather than searching for more seeds. Additionally, our field study showed that the seed encounter rate of G. allenbyi was higher than that of G. pyramidum. This difference may give G. allenbyi an advantage when resource levels are low and when searching occupies most of the forager’s time. The different advantages that each species has over the other, under different conditions, may well be factors promoting their coexistence over a wide range of resource densities. Key words: coexistence, encounter rate, foraging strategy, functional response, handling time, harvest rate. [Behav Ecol 12:219–226 (2001)]

Larger species tend to occupy more habitats, but a theoretical framework for the pattern is lacking. I modified the continuous-time logistic equation of population growth in two ways to allow for such a habitat-based theoretical framework. First, I separated birth rate from death rate. Second, I included two new terms in the equation: (1) an explicit spatial variable for habitat quality that reflects the match between a habitat and a population (species–habitat match), and (2) a demand/supply function that depends on the ratio between the energy used by all populations occurring in a habitat, and energy available in that habitat. Energy was used as a common currency to overcome differences between species of different body sizes as well as to overcome differences caused by disproportional intra- and interspecific effects. Allometric relations were used to characterize parameter values that correlate with body size, such as metabolic rate, birth rate, and death rate. The analytical solution of the equation for carrying capacity shows that, for a population to have a positive carrying capacity, its ratio of death rate to birth rate should be less than its match to the habitat it occupies. Literature-based body-size-dependent birth and death rates of Eutherian mammals show that the death-rate:birth-rate ratio decreases with body size. Combining the analytical solution and the death-rate:birth-rate ratio reveals that habitat generality should positively scale with body size. I used this model to simulate simple spatially explicit landscapes having diverse habitats and combinations of species of various body sizes. Using realistic parameters, the model generates results that are consistent with field observations. Thus, one can focus on specific processes to explore macroecological questions.

Grazing plays an important role in shaping ecological communities in human-related ecosystems. Although myriad studies have explored the joint effect of grazing and climate on plant communities, this interactive effect has rarely been studied in animals. We hypothesized that the effect of grazing on the reptile community varies along a climatic gradient in relation to the effect of grazing on habitat characteristics, and that grazing differentially affects reptiles of different biogeographic regions. We tested our hypotheses by collecting data on environmental characteristics and by trapping reptiles in four heterogeneous landscapes experiencing differing grazing intensities and distributed along a sharp climatic gradient. We found that while reptile diversity increased with grazing intensity at the mesic end of the gradient, it decreased with grazing intensity at the arid end. Moreover, the proportion of reptile species of differing biogeographic origins varied with the interactive effect of climate and grazing. The representation of species originating in arid biogeographic zones was highest at the arid end of the climatic gradient, and representation increased with grazing intensity within this area. Regardless of the climatic context, increased grazing pressure results in a reduction in vegetation cover and thus in changes in habitat characteristics. By reducing vegetation cover, grazing increased habitat heterogeneity in the dense mesic sites and decreased habitat heterogeneity in the arid sites. Thus, our results suggest that the same direction of habitat alteration caused by grazing may have opposite effects on biodiversity and community composition in different climatic contexts.

The yucca-yucca moth interaction is a classic case of obligate mutualism. Female moths pollinate and oviposit in the gynoecium of the flower; however, maturing larvae eat a fraction of the developing seeds. We studied within-fruit distributions of four seed types (fertile, infertile, eaten and uneaten seeds) in order to evaluate costs and benefits in aYucca schottii population in southeastern Arizona. We focused on how the spatial arrangement of seeds affected larval behaviour and, hence, the costs of the mutualism to the yucca. Infertile seeds were distributed throughout both infested and uninfested locules. Additionally, moth larvae feeding in a single locule preferred fertile seeds and even avoided infertile seeds and left the fruit significantly more often when they encountered infertile seeds. We suggest that, regardless of the cause of infertile seeds, they function as blocking units within seed locules and therefore reduce seed predation by moth larvae. We also suggest that, together with certain other fruit traits, the presence of infertile seeds promotes the evolutionary stability of this pollination mutualism.

Quantifying the association of plant functional traits to environmental gradients is a promising approach for understanding and projecting community responses to land use and climatic changes. Although habitat fragmentation and climate are expected to affect plant communities interactively, there is a lack of empirical studies addressing trait associations to fragmentation in different climatic regimes.In this study, we analyse data on the key functional traits: specific leaf area (SLA), plant height, seed mass and seed number. First, we assess the evidence for the community assembly mechanisms habitat filtering and competition at different spatial scales, using several null-models and a comprehensive set of community-level trait convergence and divergence indices. Second, we analyse the association of community-mean traits with patch area and connectivity along a south–north productivity gradient.We found clear evidence for trait convergence due to habitat filtering. In contrast, the evidence for trait divergence due to competition fundamentally depended on the null-model used. When the null-model controlled for habitat filtering, there was only evidence for trait divergence at the smallest sampling scale (0.25 m × 0.25 m). All traits varied significantly along the S–N productivity gradient. While plant height and SLA were consistently associated with fragmentation, the association of seed mass and seed number with fragmentation changed along the S–N gradient.Our findings indicate trait convergence due to drought stress in the arid sites and due to higher productivity in the mesic sites. The association of plant traits to fragmentation is likely driven by increased colonization ability in small and/or isolated patches (plant height, seed number) or increased persistence ability in isolated patches (seed mass).Our study provides the first empirical test of trait associations with fragmentation along a productivity gradient. We conclude that it is crucial to study the interactive effects of different ecological drivers on plant functional traits.

The response of species diversity to dispersal capability is inherently scale‐dependent: increasing dispersal capability is expected to increase diversity at the local scale, while decreasing diversity at the metacommunity scale. However, these expectations are based on model formulations that neglect dispersal limitation and species segregation at the local scale. We developed a unifying framework of dispersal–diversity relationships and tested the generality of these expectations. For this purpose we used a spatially‐explicit neutral model with various combinations of survey area (local scale) and landscape size (metacommunity scale). Simulations were conducted using landscapes of finite and of conceptually infinite size. We analyzed the scale‐dependence of dispersal‐diversity relationships for exponentially‐bounded versus fat‐tailed dispersal kernels, several levels of speciation rate and contrasting assumptions on recruitment at short dispersal distances. We found that the ratio of survey area to landscape size is a major determinant of dispersal–diversity relationships. With increasing survey‐to‐landscape area ratio the dispersal–diversity relationship switches from monotonically increasing through a U‐shaped pattern (with a local minimum) to a monotonically decreasing pattern. Therefore, we provide a continuous set of dispersal–diversity relationships, which contains the response shapes reported previously as extreme cases. We suggest the mean dispersal distance with the minimum of species diversity (minimizing dispersal distance) for a certain scenario as a key characteristic of dispersal–diversity relationships. We show that not only increasing mean dispersal distances, but also increasing variances of dispersal can enhance diversity at the local scale, given a diverse species pool at the metacommunity scale. In conclusion, the response of diversity to variations of dispersal capability at spatial scales of interest, e.g. conservation areas, can differ more widely than expected previously. Therefore, land use and conservation activities, which manipulate dispersal capability, need to consider the landscape context and potential species pools carefully.

Patterns of genetic differentiation among taxa at early stages of divergence provide an opportunity to make inferences about the history of speciation. Here, we conduct a survey of DNA-sequence polymorphism and divergence at loci on the autosomes, X chromosome, Y chromosome and mitochondrial DNA in samples of Mus domesticus, M. musculus and M. castaneus. We analyzed our data under a divergence with gene flow model and estimate that the effective population size of M. castaneus is 200 000–400 000, of M. domesticus is 100 000–200 000 and of M. musculus is 60 000–120 000. These data also suggest that these species started to diverge approximately 500 000 years ago. Consistent with this recent divergence, we observed considerable variation in the genealogical patterns among loci. For some loci, all alleles within each species formed a monophyletic group, while at other loci, species were intermingled on the phylogeny of alleles. This intermingling probably reflects both incomplete lineage sorting and gene flow after divergence. Likelihood ratio tests rejected a strict allopatric model with no gene flow in comparisons between each pair of species. Gene flow was asymmetric: no gene flow was detected into M. domesticus, while significant gene flow was detected into both M. castaneus and M. musculus. Finally, most of the gene flow occurred at autosomal loci, resulting in a significantly higher ratio of fixed differences to polymorphisms at the X and Y chromosomes relative to autosomes in some comparisons, or just the X chromosome in others, emphasizing the important role of the sex chromosomes in general and the X chromosome in particular in speciation.

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.

Conservation biologists worry that fragmenting a bloc of natural habitat might reduce its species diversity. However, they also recognize the difficulty and importance of isolating the effect of fragmentation from that of simple loss of area. Using two different methods (species–area curve and Fisher's α index of diversity) to analyse the species diversities of plants, tenebrionid beetles and carabid beetles in a highly fragmented Mediterranean scrub landscape, we decoupled the effect of degree of fragmentation from that of area loss. In this system, fragmentation by itself seems not to have influenced the number of species. Our results, obtained at the scale of hectares, agree with similar results at island and continent scales.

Mutualisms often involve significant costs for participants. Costs are inflicted by mutualists themselves, as well as by associated, non-mutualistic species. These costs are rarely quantified, however, particularly the ones extrinsic to the pairwise interaction. We compare costs inflicted by an obligate mutualist pollinator and two common exploiters of an Arizona yucca over a 2-year period. The magnitude of seed damage from seed and fruit-feeding beetle larvae (Carpophilus longus, Nitidulidae) was similar to damage from the seed-eating larvae of Yucca schottii's pollinator moth Tegeticula yuccasella (Prodoxidae), averaging about 15 seeds destroyed per fruit in each case. The two seed predators usually fed within the same fruits, although rarely side by side. In contrast, the presence of fruit-galling moth larvae (Prodoxusy-inversus, Prodoxidae) appeared to benefit the yucca: individual Tegeticula destroyed only half as many seeds in galled fruits as they did in ungalled fruits. We discuss three general implications of these results. Firstly, the costs of non-mutualists to the two mutualistic partners are not necessarily parallel. Secondly, measurable costs of non-mutualists do not necessarily translate into an impact on the success of the mutualism itself, because they may be incurred after mutualistic activities take place. Finally, the costs of mutualists to each other can differ substantially depending on the presence or absence of non-mutualistic species.

Two coexisting gerbil species, Gerbillus allenbyi (26 g) andG. pyramidum (40 g), exhibit shared-preference optimal habitat selection. In a mosaic of stabilized sands and semistabilized dunes, both species primarily prefer the semistabilized-dune habitat. We hypothesized that foraging benefit, due to the lower amount of loess in the semistabilized dunes, is the underlying cause for this preference. We tested this by measuring foraging efficiencies (giving-up density of seeds) in both manipulated foraging substrates (sand and loess) and natural habitats (semistabilized dunes, stabilized sands, and loess plateau). We also compared harvest rates on sand and loess. We used the loess substrate and loess plateau habitat to exaggerate the effect of loess on the gerbils' foraging behavior. Regarding foraging substrate, gerbils had significantly higher foraging efficiencies on sand in the field. They also had significantly higher harvest rates on sand in the laboratory. Regarding habitats, both species were more efficient in semistabilized dunes than in stabilized sands or loess plateau. Additional results suggested that the area surrounding a foraging patch had an important role in predator avoidance. We concluded that the semistabilized-dune habitat allowed gerbils to forage more efficiently in a patch, because it is easier to dig for seeds in the sandy substrate found there, and because gerbils enjoyed lower predatory risk traveling to and from the seed patch. The results supported the hypothesis that foraging benefit is the basis of the shared-preference habitat selection of the gerbil species

We have tested two hypotheses which may explain two different patterns which underlie coexistence in two species of desert gerbils (Gerbillus allenbyi and G. pyramidum). The two patterns are temporal partitioning of foraging activity and shared preference habitat selection. When sympatric, G. pyramidum uses the early part of the night most heavily while G. allenbyi is most active in the late part of the night. Although both species prefer the same habitat type (semistabilized sand dune), in the presence of G, pyramidum, G. allenbyi utilizes mainly the secondary habitat type (stabilized sand dune)