Body mass variations in disturbed mallards Anas platyrhynchos fit to the mass‐dependent starvation‐predation risk trade‐off

For passerines the starvation‐predation risk theory predicts that birds should decrease their body mass to improve escape flight performance, when predation pressure increases. To investigate whether this theory may apply to large birds, which manage body reserves differently from small passerines, we experimentally increased the predation risk in mallards Anas platyrhynchos. Two groups were disturbed at different frequencies during experimental sessions lasting one week, while a control group was left undisturbed. We found that body mass loss and final wing loading were similar in both disturbed groups and significantly differed from the control group. Food intake in disturbed groups was reduced up to day four of the disturbance session and was lower than in the control group. Altogether our results suggest that disturbed mallards may adjust their body mass to reach a more favorable wing loading, supposedly to improve escape flight performance. Nevertheless, body mass loss in our mallards was double than what has been observed in passerines. This greater mass decrease might be explained by different strategies concerning energy storage. Furthermore, in large birds the predation component of the starvation‐predation trade‐off might be of greater importance. Hence, the observed relevance of this trade‐off over a large size range suggests that the starvation‐predation risk theory is of major ecological significance for many animal species.     

Management of reedbeds: mosaic reed cutting does not affect prey abundance and nest predation rate of reed passerine birds

Reed passerine birds are strict habitat specialists inhabiting reedbed habitats. In Europe, many of these species are threatened due to loss and degradation of natural reedbeds. Another important factor that can negatively affect the abundance of reed passerines is commercial reed harvesting. Previous studies have shown negative impacts of large-scale winter reed cutting on passerine breeding assemblages and arthropod communities. The effect of reed cutting on a small scale, however, has not been studied experimentally to date. The aim of this study was to investigate whether and how small-scale, mosaic reed cutting influences prey abundance and nest predation rate of reed passerines. In June, after the reed had reached maturity, we conducted nest predation experiments with artificial nests and arthropod sampling using pan traps in cut reed patches, adjacent uncut reed patches and unmanaged reedbed. We found no differences in the risk of egg predation between three types of reedbeds. In contrast, the abundance of arthropods in cut and adjacent uncut reed patches was significantly higher than that in unmanaged reedbed. We assume this was caused by habitat heterogeneity, small size of cut patches and their rapid recolonization by arthropods from adjacent uncut patches. Our results suggest that in contrast to large-scale reed cutting, small-scale, mosaic reed cutting has no negative effect on nest survival and food abundance of reed passerine birds. However, given that we performed all experiments in June, i.e., when the reed was mature, our findings cannot be generalized to whole breeding period of all reed passerine birds. Therefore, temporal variation in nest predation rate and arthropod abundance in managed and unmanaged reedbeds during the entire breeding season should be examined in future studies.     

捕食风险对不同饥饿状态下艾虎取食行为的影响

在室内条件下,将大鵟作为艾虎的天敌动物,通过双通道选择实验确定6 只成体艾虎在3 个捕食风险水平和4 种饥饿状态条件下的取食行为,探讨艾虎在取食过程中对饥饿风险与捕食风险的权衡策略。研究结果表明:在无捕食风险存在时,艾虎被剥夺食物0 d 和1 d 后对食物量不同的两个斑块中的取食量和利用频次均无明显不同(P > 0. 05),但对高食物量斑块的利用时间均明显高于低食物量斑块的(P <0.05),而艾虎被剥夺食物2 d和3 d后对高食物量斑块中的取食量和利用时间均明显高于低食物量斑块中的(P < 0.05),但在利用频次上均无明显差异(P > 0.05)。在面临低风险时,艾虎在4 种饥饿状态下均只利用无天敌动物存在的低食物量斑块,而基本不利用有天敌动物存在的高食物量斑块。在面临高风险时,艾虎不得不利用有天敌动物存在的食物斑块,被剥夺食物0 d 时艾虎对无风险、无食物量斑块的利用时间基本相同于对高风险、有食物量斑块的利用时间(P>0.05),而被剥夺食物1d、2 d 和3 d 后艾虎对高风险、有食物量斑块的利用时间明显高于无风险、无食物量斑块的(P< 0. 05)。在相同风险条件下,随着饥饿程度增加,艾虎在斑块中的取食量均明显增加(P< 0.05),而对斑块的利用时间和利用频次明显降低(P<0.05)。在相同的饥饿状态下,不同风险水平时,艾虎在斑块中的取食量无明显的差异(P>0.05),但在低风险和高风险时对斑块的利用时间和频次均明显低于无风险时的(P <0.05)。以上结果说明艾虎能够根据食物摄取率和自身的能量需求在捕食风险和饥饿风险之间做出权衡,当饥饿风险小于捕食风险时,艾虎趋于躲避捕食风险,当饥饿风险大于捕食风险时,艾虎趋于面对捕食风险,所采用的取食策略是减少活动时间和能量消耗,最大程度地提高单位时间内获得的能量。     

Within- and between-patch variability of predation intensity on the mussel Mytilus trossulus Gould on a rocky intertidal shore in Oregon, USA

Variability of predation intensity is an important cause of spatial differences of community structure and organization in the intertidal rocky shore. Field experiments were conducted to evaluate the within- and between-patch variability of the effects of different types of predators (small invertebrates and birds) on Mytilus trossulus Gould, which occupies an intermediate position in the competitive hierarchy among sessile organisms in disturbance patches within a California mussel (Mytilus californianus Conrad) bed community on the central Oregon Coast. Predation by birds did not significantly affect the mortality of M. trossulus. On the contrary, predation by small invertebrate whelks (Nucella spp.) had a significant effect on M. trossulus mortality. Predation by whelks also caused between- and within-patch variability of mortality of M. trossulus. Within patches, M. trossulus mortality at patch margins was significantly higher than at patch centers only when invertebrate predators were present. Wave exposure did not cause between-patch variability of predation intensity. Between-and within-patch variability of predation intensity may be caused by the variability of supply of whelks from the surrounding mussel mat. The movement of predators between patches and surrounding matrices may play an important role in the patch dynamics of these communities.     

Parasites and Scottish ptarmigan numbers

Summary Unlike red grouse (Lagopus lagopus scoticus), which have almost 100% prevalence of the parasitic threadworm Trichostrongylus tenuis and frequently high tapeworm numbers, 70% (n=71) of Scottish ptarmigan (Lagopus mutus) had no threadworms and only 4% had tapeworms. Other parasites and pathogenic bacteria were infrequent. Threadworms occurred in 12% (n=25) of birds on granite hills where mean ptarmigan densities were low, and in 43% (n=46) of birds over schists and limestones where mean densities were high. The notion that parasites cause the cyclic-type ptarmigan declines observed on the granite Cairngorms massif is unlikely (ptarmigan fluctuations over the richer rocks are irregular).     

High brood patch temperature of less colourful,less pheomelanic female Barn Swallows Hirundo rustica

In birds, even a minor difference in egg temperature (1–1.5 °C) has been shown to affect the fitness of offspring by changing hatching success, incubation period and nestling quality. Female, but not male, passerines develop brood patches. Thus if there are traits, such as plumage ornamentation, that indicate optimal egg temperature, males should pair with females that exhibit those traits. However, no study has yet investigated the relationship between female brood patch temperature, which would directly affect egg temperature, and female plumage ornamentation. In this study, we examined the surface temperature of female brood patches during nocturnal incubation and examined its relationship with female plumage ornaments in Asian Barn Swallows Hirundo rustica gutturalis. After controlling for ambient air temperature, brood patch temperature was negatively associated with colour saturation of the female throat patch. No other female ornaments, such as tail‐length, white tail spots or throat patch size, predicted brood patch temperature. When oral (mouth) temperature was statistically controlled, females with less colourful throats and longer tails showed higher brood patch temperature, indicating that these females had hotter brood patches in relation to the temperature of other body parts. Furthermore, we found a negative relationship between pheomelanin pigmentation and brood patch temperature after controlling for ambient air temperature or oral temperature. To our knowledge, this is the first study to show that female ornaments can predict the absolute/relative thermal investment in brood patches. This relationship, together with other aspects of female quality, may affect male mate preference and female ornamentation.     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

The Little Miss Muffet Effect: Quantifying the Effect of Predation Risk on Foraging Patch Choice by Houseflies (<Emphasis Type="Italic">Musca domestica</Emphasis>)

The ability of the ideal free distribution (IFD) to predict patch choice of female houseflies (Musca domestica) was determined by examining their distribution between two patches containing unequal amounts of food. The effect of predation risk was then quantified in energetic terms by examining fly distribution between patches of equal food, with one containing spiders. Results were used to predict how much extra food must be added to the risky patch to offset the risk of predation. Flies were found to conform fairly closely to the IFD. Predation risk had a major effect on their distribution, with fewer flies feeding in the presence of predators as risk increased. Addition of extra food to the risky patch was successful in offsetting the risk of predation. These results suggest that the effect of risk on housefly foraging behavior can be quantified in energy terms, providing a common currency for predicting the effects of resources and predation risk on habitat use.     

Behavioural Response of Juvenile Bluegill Sunfish to Variation in Predation Risk and Food Level

The behavioural response of juvenile bluegill sunfish (Lepomis macrochirus) to predation risk when selecting between patches of artificial vegetation differing in food and stem density was investigated. Bluegill foraging activity was significantly affected by all three factors. Regardless of patch stem density or risk of predation bluegills preferred patches with the highest prey number. During each trial bluegill foraging activity was clearly divided into a between- and within-patch component. In the presence of a predator bluegills reduced their between-patch foraging activity by an equivalent amount regardless of patch stem density or food level, apparently showing a risk-adjusting behavioural response to predation risk. Within patches, however, foraging activity was affected by both food level and patch stem density. When foraging in a patch offering a refuge from predation, the presence of a predator had no effect on bluegill foraging activity within this patch. However, if foraging in a patch with only limited refuge potential, bluegill foraging activity was reduced significantly in the presence of a predator. Further, this reduction was significantly greater if the patch contained a low versus a high food level, indicating a risk-balancing response to predation with respect to within-patch foraging activity. Both these responses differ from the risk-avoidance response to predation demonstrated by juvenile bluegills when selecting among habitats. Therefore, our results demonstrate the flexibility of juvenile bluegill foraging behaviour.     

Patch use in time and space for a meso-predator in a risky world

Predator–prey studies often assume a three trophic level system where predators forage free from any risk of predation. Since meso-predators themselves are also prospective prey, they too need to trade-off between food and safety. We applied foraging theory to study patch use and habitat selection by a meso-predator, the red fox. We present evidence that foxes use a quitting harvest rate rule when deciding whether or not to abandon a foraging patch, and experience diminishing returns when foraging from a depletable food patch. Furthermore, our data suggest that patch use decisions of red foxes are influenced not just by the availability of food, but also by their perceived risk of predation. Fox behavior was affected by moonlight, with foxes depleting food resources more thoroughly (lower giving-up density) on darker nights compared to moonlit nights. Foxes reduced risk from hyenas by being more active where and when hyena activity was low. While hyenas were least active during moon, and most active during full moon nights, the reverse was true for foxes. Foxes showed twice as much activity during new moon compared to full moon nights, suggesting different costs of predation. Interestingly, resources in patches with cues of another predator (scat of wolf) were depleted to significantly lower levels compared to patches without. Our results emphasize the need for considering risk of predation for intermediate predators, and also shows how patch use theory and experimental food patches can be used for a predator. Taken together, these results may help us better understand trophic interactions.     

Does tiger shark predation risk influence foraging habitat use by bottlenose dolphins at multiple spatial scales?

Prey availability and predation risk are important determinants of habitat use, but their importance may vary across spatial scales. In many marine systems, predator and prey distributions covary at large spatial scales, but do no coincide at small spatial scales. We investigated the influences of prey abundance and tiger shark ( Galeocerdo cuvier ) predation risk on Indian Ocean bottlenose dolphin ( Tursiops aduncus ) habitat use across multiple spatial scales, in Shark Bay, Western Australia. Dolphins were distributed between deep and shallow habitats and across microhabitats within patches approximately proportional to prey density when shark abundance was low. When shark abundance was high, foraging dolphins greatly reduced their use of dangerous, but productive, shallow patches relative to safer deep ones. Also, dolphins reduced their use of interior portions of shallow patches relative to their edges, which have higher predator density but lower intrinsic risk (i.e. a higher probability of escape in an encounter situation). These results suggest that predation risk and prey availability influence dolphin habitat use at multiple spatial scales, but intrinsic habitat risk, and not just predator encounter rate, is important in shaping dolphin space use decisions. Therefore, studies of habitat use at multiple spatial scales can benefit from integrating data on prey availability and the subcomponents of predation risk.     

Patch Size and Tree Species Influence the Number and Duration of Bird Visits in Forest Restoration Plots in Southern Costa Rica

Efforts to restore tropical forest in abandoned pasture are likely to be more successful when bird visitation is promoted because birds disperse seeds and eat herbivorous arthropods that damage leaves. Thus, it is critical to understand bird behavior in relation to different restoration strategies. We measured the likelihood of visitation, number of visits, and duration of visits for all birds and for Cherrie's Tanager ( Ramphocelus costaricensis ), a common seed disperser, in five sizes of forest restoration patches planted with four tree species in southern Costa Rica. We predicted that the largest patches, and the tree species with the greatest canopy cover, would be visited most frequently and have the longest visits because we assumed that these patch types had the greatest food resources and the lowest predation risk. We found that birds were more likely to visit large patches and the tree species with the highest canopy cover ( Inga edulis ). Birds visited Inga trees more often and stayed in Inga and Erythrina poeppigiana trees for longer periods of time than in other tree species. We found similar results for Cherrie's Tanagers. Thus, we identified two factors, tree species and patch size, which may be manipulated in restoration projects to increase bird visitation.     

Comparative isotopic natural history of two native passerines (Troglodytes cobbi and Cinclodes antarcticus) and the invasive rats (Rattus norvegicus) that extirpate them

While several studies have shown that invasive rats can have negative effects on island birds through predation (both direct predation and nest predation), other mechanisms for the effects of invasives on island biota have been given less attention. Here, we explore another potential mechanism by which invasive rats can affect native island birds: the competitive use of common resources. We used stable isotope analyses to estimate the fraction of marine and terrestrial sources incorporated into the tissues of two species of passerines (Troglodytes cobbi, Troglodytidae; and Cinclodes antarcticus, Furnariidae) and Norway rats (Rattus norvegicus, Muridae) in the Falkland Islands. These two passerines are absent on islands where rats are present. We found significant incorporation of marine resources in the three species, with the highest incorporation in tissues of T. cobbi. This species appears to be one of the passerines most reliant on marine sources and the most marine member of the family Troglodytidae. We also used the results of these isotopic analyses to estimate the isotopic niche breadth of each of these species and the isotopic niche overlap among them. Rattus norvegicus had a large isotopic niche that overlapped broadly with those of the two passerine species. We propose that different ways of both depicting and estimating isotopic niche widths are complementary rather than alternative. Our results are consistent with the notion that invasive rats might have an impact on these two species of Falkland Island passerines by using common resources but do not rule out the possibility that part of their effect is through direct predation.     

Evidence of the trade-off between starvation and predation risks in ducks

The theory of trade-off between starvation and predation risks predicts a decrease in body mass in order to improve flight performance when facing high predation risk. To date, this trade-off has mainly been validated in passerines, birds that store limited body reserves for short-term use. In the largest avian species in which the trade-off has been investigated (the mallard, Anas platyrhynchos), the slope of the relationship between mass and flight performance was steeper in proportion to lean body mass than in passerines. In order to verify whether the same case can be applied to other birds with large body reserves, we analyzed the response to this trade-off in two other duck species, the common teal (Anas crecca) and the tufted duck (Aythya fuligula). Predation risk was simulated by disturbing birds. Ducks within disturbed groups were compared to non-disturbed control birds. In disturbed groups, both species showed a much greater decrease in food intake and body mass during the period of simulated high risk than those observed in the control group. This loss of body mass allows reaching a more favourable wing loading and increases power for flight, hence enhancing flight performances and reducing predation risk. Moreover, body mass loss and power margin gain in both species were higher than in passerines, as observed in mallards. Our results suggest that the starvation-predation risk trade-off is one of the major life history traits underlying body mass adjustments, and these findings can be generalized to all birds facing predation. Additionally, the response magnitude seems to be influenced by the strategy of body reserve management.     

Seasonal variation in patch use in a tropical African environment

We used giving-up densities (GUD) to study patch use decisions of small granivorous passerines throughout the year. We measured GUDs continuously in four sites for a period of 9–10 months per year during 2004 and 2005 in a savannah area in Jos, central Nigeria. The study thus covered a period from the middle of the dry season, through the wet season to the beginning of the next dry season in each year. We placed experimental food patches in both open areas and in cover to investigate possible effects of predation risk and thermal hazard on the foraging behavior of the birds. We found a difference in GUDs between the microhabitats, with a consistently lower GUD in cover throughout the year and for the two years. During both years GUDs followed a pattern coinciding with the seasonal change in local seed availability. An initial decline in GUDs late in the dry season was followed by a steady increase during and after the rains. A similar trend in GUDs observed for both years supports the conclusion that GUDs measure the feeding birds' assessment of environmental quality, possibly in combination with other factors changing predictably during the year. We conclude that food abundance may act with other environmental and ecological factors to affect foraging decisions throughout the year.     

Leaving final-cut grass silage in situ overwinter as a seed resource for declining farmland birds

The loss of seed-rich wintering habitats has been a major contributory cause of farmland bird population declines in western Europe. Agricultural grasslands are particularly poor winter foraging habitats for granivorous birds, which have declined most in the pastoral farming regions of western Britain. We describe an experiment to test the utility of fertile ryegrass (Lolium) swards as a potentially rich source of winter seed for declining farmland birds. Four patches of final-cut grass silage were allowed to set seed and were left in situ overwinter. Half of each patch was lightly aftermath grazed in an attempt to increase the accessibility of the seed to foraging birds and reduce the perceived predation risk. Large numbers of yellowhammers (Emberiza citrinella) and reed buntings (E. schoeniclus) foraged on the seeded plots throughout the winter. They preferred to forage on ungrazed seeded plots, where the accumulation of senescent foliage resulted in a 14% average loss in silage yield in the following season. However, seed produced on the plots also led to sward regeneration, increasing subsequent yields on some plots. The technique offers clear benefits as a potential future agri-environment measure for declining granivorous birds, with wide applicability, but requires further development to minimise sward damage and costs to the farmer. Autumn grazing should reduce sward damage, but at the cost of reduced usage by buntings. Using the technique just prior to reseeding would be one way of avoiding any costs of sward damage.     

The Use and Transfer of Information About Predation Risk in Flocks of Wintering Finches

Several studies in behavior have focused in some way on how groups of prey gather and use information about predation risk. Although asymmetries in information about risk exist among members of real groups, we know little about how such uneven information might affect individual or group antipredator decisions. Hence, we studied the use and transfer of information about the risk of predation in small flocks of wintering birds. House finches (Carpodacus mexicanus; 28 groups of three) were held in large enclosures divided into safe and risky patches. We controlled the information about risk available to each individual by conducting attacks with a model hawk that was visible to only a single (informed) bird. Repeated attacks on a single individual did not reduce the amount of feeding by other birds in that patch, although the time to resume feeding after observing a response to an attack event was somewhat longer than after a no attack event. These results suggest that informed individuals impart some information to naïve (uninformed) birds, but this effect was not strong. In fact, the frequent return of informed individuals to feeders after observing uninformed individuals feed suggests that finches relied more on public information regarding safety than their own personal information in deciding when to feed. Group patch choice appeared to be based on a majority‐rules decision, although an effect of dominance status was apparent. Our results suggest that subordinate flock members may exert a large influence over group decision‐making by acting as spatial ‘anchors’.     

Titrating the cost of plant toxins against predators: determining the tipping point for foraging herbivores

1. Foraging herbivores must deal with plant characteristics that inhibit feeding and they must avoid being eaten. Principally, toxins limit food intake, while predation risk alters how long animals are prepared to harvest resources. Each of these factors strongly affects how herbivores use food patches, and both constraints can pose immediate proximate costs and long-term consequences to fitness. 2. Using a generalist mammalian herbivore, the common brushtail possum (Trichosurus vulpecula), our aim was to quantitatively compare the influence of plant toxin and predation risk on foraging decisions. 3. We performed a titration experiment by offering animals a choice between non-toxic food at a risky patch paired with food with one of five toxin concentrations at a safe patch. This allowed us to identify the tipping point, where the cost of toxin in the safe food patch was equivalent to the perceived predation risk in the alternative patch. 4. At low toxin concentration, animals ate more from the safe than the risky patch. As toxin concentration increased at the safe patch, intake shifted until animals ate mainly from the risky patch. This shift was associated with behavioural changes: animals spent more time and fed longer at the risky patch, while vigilance increased at both risky and safe patches. 5. Our results demonstrate that the variation in toxin concentration, which occurs intraspecifically among plants, can critically influence the relative cost of predation risk on foraging. We show that herbivores quantify, compare and balance these two different but proximate costs, altering their foraging patterns in the process. This has potential ecological and evolutionary implications for the production of plant defence compounds in relation to spatial variation in predation risk to herbivores.     

Habitat use under latent predation risk. A case study with wintering forest birds

We test whether the spatial distribution of birds within a habitat is determined by predation risk and also by interspecific competition. The work was carried out in a montane mixed forest of central Spain with four Parus species, the long-tailed tit ( Aegithalos caudatus ) and the nuthatch ( Sitta europaea ). Experimental feeders, that varied in their risk of predation, were used to control the effect of natural variations in food availability and quality on the habitat use patterns of different species. Tree gleaning passerines avoided feeding on dark inner forest places far from edges, distant from protective cover, outside the inner tree canopy and near the ground; they preferred deciduous, relatively clear forest plots. These effects remained invariable across years and weather conditions. There was a common pattern of selection of foraging locations by the four Parus species: proximity to cover and height above ground and over the lowest branches of the tree canopy positively influenced the use of feeding places. According to these patterns, the vigilance proportion of species was significantly higher when feeding far from cover than when birds were feeding near pine foliage. This pattern was also common for the four studied Parus species. Nevertheless, the interspecific dominance hierarchy of the species was positively correlated with the use of the safest feeders (feeders farther the ground and nearer from protective cover within tree canopy), being the converse with the most exposed ones. Therefore, the results of this paper demonstrate that the selection of feeding locations within habitat follows a pattern minimizing predation risk. Interspecific dominance hierarchies can lead to the exploitation of unfavourable risky patches by subordinate species.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).