Sexual segregation in ungulates: a comparative test of three hypotheses

In most social ungulate species, males are larger than females and the sexes live in separate groups outside the breeding season. It is important for our understanding of the evolution of sociality to find out why sexual segregation is so widespread not only in ungulates but also in other mammals. Sexual body size dimorphism was proposed as a central factor in the evolution of sexual segregation in ungulates. We tested three hypotheses put forward to explain sexual segregation: the predation-risk, the forage-selection, and the activity budget hypothesis. We included in our analyses ungulate species ranging from non-dimorphic to extremely dimorphic in body size. We observed oryx, zebra, bighorn sheep and ibex in the field and relied on literature data for 31 additional species. The predation-risk hypothesis predicts that females will use relatively predator-safe habitats, while males are predicted to use habitats with higher predation risk but better food quality. Out of 24 studies on different species of ungulates, females and their offspring chose poorer quality but safer habitat in only eight cases. The forage-selection hypothesis predicts that females would select habitat based on food quality, while males should prefer high forage biomass. In fact, females selected higher quality food in only six out of 18 studies where males and females segregated, in eight studies there was no difference in forage quality and in four studies males were in better quality habitat. The activity budget hypothesis predicts that with increasing dimorphism in body size males and females will increasingly differ in the time spent in different activities. Differences in activity budgets would make it difficult for males and females to stay in mixed-sex groups due to increased costs of synchrony to maintain group cohesion. The predictions of the activity budget hypothesis were confirmed in most cases (22 out of 23 studies). The heavier males were compared to females, the more time females spent foraging compared to males. The bigger the dimorphism in body mass, the more males spent time walking compared to females. Lactating females spent more time foraging than did non-lactating females or males. Whether species were mainly bulk or intermediate feeders did not affect sexual differences in time spent foraging. We conclude that sexual differences in activity budgets are most likely driving sexual segregation and that sexual differences in predation risk or forage selection are additive factors.     

Sexual segregation in vertebrates: proximate and ultimate causes

Sexual segregation is very common in vertebrates that live ingroups. In this article, I will review proximate and ultimatecauses of sexual segregation in social species and in particularin ungulates in which the bulk of research on the topic hasbeen carried out. In most social ungulate species, males andfemales live in separate groups outside the breeding season,sometimes using different home ranges and types of habitat.In most of these species, males are larger than females. Dimorphismin body size can lead to sexual differences in ecology and behaviormaking it difficult for the two sexes to stay in the same group.It is important for our better understanding of the evolutionof sociality, sexual dimorphism and different mating systemsto determine why sexual segregation is so widespread not onlyin ungulates but also in other vertebrates. In this article,I discuss the ecology of the two sexes by reviewing proximateand ultimate causes of sexual segregation. To do this, I comparea range of studies of ruminants and include explanations forsocial segregation as well as for habitat segregation by gender.This leads into a review and updates current knowledge of thephenomenon. Although I present a number of different hypotheses,I focus in particular on predation risk, forage selection andactivity budget and discuss the social-factors hypothesis. Istress that the key in solving the enigma of sexual segregationlies in clearly separating hypotheses that try to explain socialsegregation and habitat segregation, as well as in includingexperiments or model systems. To that end, I present a preliminarystudy on a test of the activity-budget hypothesis in three-spinesticklebacks and explain why I believe that shoaling fish areuseful for analysing the underlying processes and mechanismsthat lead to sexual segregation in animals. Lastly, I arguethat it is unlikely that a single factor can explain socialsegregation or habitat segregation but that a model integratingdifferent factors and different levels of segregation mightsucceed in describing proximate and ultimate causes of sexualsegregation.     

Predation Risk Drives Habitat‐Specific Sex Ratio in a Monomorphic Species,the Brown Hare (Lepus europaeus)

In dimorphic species, sexual habitat segregation is generally explained by the differences in nutritional needs or by a trade‐off between fulfilling food requirements and avoiding predation. However, it remains unclear whether predation risk is strong enough to drive the differences in habitat use between sexes as predicted by the predation sensitivity hypothesis. Here we test in a monomorphic species, the brown hare (Lepus europaeus), the prediction that abundance of the gender more sensitive to predation is higher in safer habitat. We used data on 1645 individually marked hares in western Poland during autumn–winter seasons of 1966/1967–1978/1979 to estimate sex‐specific annual survival rates. We analyzed the stomach contents of 134 foxes shot in 1965/1966–1994/1995 to evaluate fox predation on hares. Finally, we employed data on 26 790 hares live‐trapped in 1965/1966–1994/1995 to analyze hare sex ratio across habitats. We found that male annual survival rate was lower than that of females and that the predation risk by foxes on hares was lower in agricultural than forest habitat. Our finding, that males were more often trapped by nets in agricultural than the forest habitat, provides indirect evidence for the predation sensitivity hypothesis. We conclude that predation risk can be a driving force for habitat‐specific sex ratio in a monomorphic species such as the brown hare.     

Differential Migration by Sex in the Great Bustard: Possible Consequences of an Extreme Sexual Size Dimorphism

We explored migration patterns in Great bustards ( Otis tarda ), a species that shows strong sexual selection and the most extreme sexual size dimorphism among birds. The aim was to explain differential migration, examining whether Great bustards fulfil the main predictions of bird migration theory hypotheses and sexual segregation theory hypotheses. We radio-tracked the seasonal movements of 65 males and 68 females in central Spain. We found four main sexual differences. First, the proportion of migratory males was higher than that of migratory females (86% vs. 51%). Second, males abandoned the leks immediately after the mating season (late May to early Jun.), whereas females remained there for another 3–7 mo. Third, 54% of the migratory males used two different post-breeding areas, the first located northwards at 82 km from the breeding sites in summer, and the second southwards at 50 km in autumn–winter. Migratory females used only one area in autumn–winter which coincided geographically with that of males. And fourth, males returned to the breeding areas earlier (between Sep. and Mar.) than females (between Jan. and Apr.). These results show that the Great bustard is a differential migrant by sex in central Spain and support the weather sensitivity hypothesis (males were less tolerant to summer heat) and the specialization hypothesis (exclusive maternal care of the brood by females). Sexual differences in migratory behaviour are probably ultimately determined by the strong sexual selection in this species.     

Is sexual segregation in the guppy, <Emphasis Type="Italic">Poecilia reticulata</Emphasis>, consistentwith the predation risk hypothesis?

We examined the importance of sex differences in predation risk in generating sexual segregation in the guppy, Poecilia reticulata. We hypothesised that sex differences in predation risk will result in habitat segregation and ultimately social segregation of the sexes, with the more vulnerable sex (males in this case) using safer habitats. In accordance with the predation risk hypothesis we observed sexual segregation in a population associated with high but not low predation risk. Under high predation risk we observed a larger proportion of males in shallow marginal habitats resulting in habitat segregation and ultimately social segregation of the sexes. Furthermore, habitat segregation by sex was associated with habitat segregation by body length with shoals in deeper water having a larger mean body length. Shoaling fish species have been key models in investigating group living, and further research directed towards understanding sexual segregation in other fish species would be valuable.     

Sex differences in weather sensitivity can cause habitat segregation: red deer as an example

Sex differences in habitat use (habitat segregation) are widespread in sexually dimorphic ungulates. A possible cause is that males are more sensitive to weather than females, leading to sex differences in sheltering behaviour (the 'weather sensitivity hypothesis'). However, this hypothesis has never been tested. We considered the allometric rates of net energy gain during times of cold weather and food shortage in a model. We argue that the higher absolute heat losses relative to intake rates of larger ungulates should indeed lead to higher weather sensitivity in males than in females. Furthermore, we tested the weather sensitivity hypothesis empirically in red deer, Cervus elaphus, on the Isle of Rum, U.K. We predicted that (1) use of relatively exposed, high-quality forage habitat should be negatively influenced by bad weather; and (2) this influence should be stronger in males. We found that bad weather (strong wind, low temperature, heavy rain) in winter and spring influenced use of high-quality forage habitat negatively in all deer; that adult males responded more strongly to low temperature and strong wind than did females; and that adult males foraged on windy days at better sheltered sites than did females. Thus, the weather sensitivity hypothesis is supported both theoretically and empirically. We suggest that the weather sensitivity hypothesis can potentially explain winter habitat segregation in a large number of ungulate species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Test of the activity budget hypothesis on Asiatic ibex in Tian Shan Mountains of Xinjiang,China

Asiatic ibex (Capra sibirica) is a threatened species in China and is distributed in the mountains of Central Asia. It is a sexually dimorphic ungulate. The males are much larger than females, and except for the breeding season, both sexes live in separate groups. Many hypotheses have been developed to explain the sexual segregation in sexually dimorphic ungulates. These hypotheses are not mutually exclusive, and in recent years, the activity budget hypothesis has received special attention. To test this hypothesis, we studied the activity budget of Asiatic ibex in the autumn of 2005. According to the activity budget hypothesis, females should spend more time feeding than the males, and the degree of activity synchronization should be higher in same-sex groups than in mixed-sex groups. Our results are consistent with the hypothesis. Both sexes of the Asiatic ibex spent most of the time feeding, and females spent a significantly longer time feeding than males, and males spent a significantly longer time standing. Activity synchronization indexes of both the female groups and males groups were significantly higher than mixed-sex groups. These results indicated that in Asiatic ibex, the activity budget hypothesis about sexual segregation is supported.     

An experimental demonstration of the impact of predation on sexual segregation and primary sex ratios among ungulates

Underlying mechanisms of sexual segregation among ungulates, and Trivers and Willard's hypothesis that mothers can influence primary sex ratios, continue to be topical theoretical issues. Over 2 years, using monthly repeated road transects, we determined the habitat and social segregation of male vs. female impala (Aepyceros melampus) and kudu (Tragelaphus strepsiceros) in a predator‐free, vs. a predator‐laden, South African reserve. We also determined, by the same technique but over 4 years, the primary sex ratio of the impala population free from predation. Significant overlap in habitat usage (Schoener's Index 0.63–0.8) was found between the two sexes when free from predation, but not (Schoener's Index 0.46–0.47) when under predation. While occupying the same habitats impala, kudu and wildebeest (Connochaetes taurinus) male and female groups maintained rigid social segregation throughout the year, even when at close quarters. Impala primary sex ratios were significantly biased towards females (male/female = 0.72; χ² = 4.3175, d.f. = 1, P‐value = 0.038) in the absence of predation. Our findings suggest that while risk of predation is a proximal cause of sexual segregation, thus lending support to the predator‐risk hypothesis, the underlying, functional mechanism of sexual segregation is the difference in the activity budgets of males vs. females (the activity‐budget hypothesis). Our findings also suggest that mothers may indeed be able to adjust primary sex ratios, with the postulated driver in this case being an abnormally high density of adult males.     

Sexual segregation in western grey kangaroos: testing alternative evolutionary hypotheses

In sexually dimorphic ungulates, sexual segregation is hypothesized to have evolved because of sex-specific differences in body size and/or reproductive strategies. We tested these alternative hypotheses in kangaroos, which are ecological analogues of ungulates. Kangaroos exhibit a wide range of body sizes, particularly among mature males, and so the effects of body size and sex can be distinguished. We tested predictions derived from these hypotheses by comparing the distribution of three sex–sex size classes of western grey kangaroos Macropus fuliginosus , in different habitats, and the composition of groups of kangaroos, across seasons. In accordance with the predation risk-reproductive strategy hypothesis, during the non-breeding season, females, which were more susceptible to predation than larger males, and were accompanied by vulnerable young-at-foot, were over-represented in secure habitats. Large males, which were essentially immune to predation, occurred more often than expected in nutrient-rich habitat, and small males, which faced competing demands of predator avoidance and feeding, were intermediate between females and large males in their distribution across habitats. During the breeding season, females continued to be over-represented in secure habitats when their newly emerged pouch young were most vulnerable to predation. All males occupied these same habitats to maximize their chances of securing mates. Consistent with the social hypotheses, groups composed of individuals of the same sex, irrespective of body size, were over-represented in the population during the non-breeding season, while during the breeding season all males sought females so that mixed-sex groups predominated. These results indicate that body size and reproductive strategies are both important, yet independent, factors influencing segregation in western grey kangaroos.     

Mechanisms of species coexistence: a field test of theoretical models using intertidal snails

Competitor coexistence is often facilitated by spatial segregation. Traditionally, spatial segregation is predicted to occur when species differ in the habitat in which they are either superior at competing for resources or less susceptible to predation. However, predictions from a behavioural model demonstrate that spatial segregation and coexistence can also occur in the absence of such interspecific trade‐offs in competitive ability and vulnerability to predation. Unlike other models of competitor coexistence this model predicts that when species rank both habitat productivity and ‘riskinesses’ similarly, but differ slightly in their habitat‐specific vulnerabilities to predators, they will tend to segregate across habitats, with the species experiencing the higher ratio of mortality risk across the habitats occurring primarily in the safer habitat. Here, we investigate the hypothesis that intraspecific trade‐offs between resource availability and mortality risk can lead to spatial segregation of competing species by (1) documenting the spatial (i.e. intertidal) distribution of two marine snails, Littorina sitkana and L. subrotundata and (2) performing field experiments to quantify growth and mortality rates of each species at ‘low’ and ‘high’ intertidal heights. Our results indicate that both species agree on the rankings of habitat riskiness and productivity, experiencing higher predation and higher growth in low‐ than in high‐intertidal habitats. However, L. sitkana and L. subrotundata experienced differences in their habitat‐specific mortality risks and growth rates. Despite both species being similarly at risk of predation in high‐intertidal habitats (where mortality was lower), L. subrotundata was subject to significantly higher mortality than L. sitkana at the low‐intertidal height. In contrast, growth rate differences between habitats were greater for L. sitkana than for L. subrotundata. Whereas both species grew at the same rate at the high‐intertidal level (where growth was lower), L. sitkana individuals grew more rapidly than L. subrotundata snails at the low‐intertidal level. As predicted by the behavioural model, the species that experienced the higher ratio of mortality across habitats (i.e. L. subrotundata) occurred exclusively in the safer, high‐intertidal habitat. Taken together, these results provide support for the hypothesis that spatial segregation, and potentially competitor coexistence, can occur in the absence of interspecific trade‐offs in resource acquisition ability or vulnerability to predation.     

The evolution of movements and behaviour at boundaries in different landscapes: a common arena experiment with butterflies

As landscapes change, mobility patterns of species may alter. Different mechanistic scenarios may, however, lead to particular patterns. Here, we tested conflicting predictions from two hypotheses on butterfly movements in relation to habitat fragmentation. According to the resource distribution hypothesis, butterflies in more fragmented landscapes would have higher levels of mobility as resources are more scattered. However, these butterflies could have lower levels of mobility as they experience 'hard' habitat boundaries more frequently (i.e. higher crossing costs) compared with butterflies in landscapes with continuous habitat; i.e. the behaviour-at-boundaries hypothesis. We studied movements, habitat boundary crossing and habitat preference of laboratory-reared individuals of Pararge aegeria that originated from woodland and agricultural landscapes, by using an experimental landscape as a common environment (outdoor cages) to test the predictions, taking into account sexual differences and weather. Woodland butterflies covered longer distances, were more prone to cross open-shade boundaries, travelled more frequently between woodland parts of the cages and were more at flight than agricultural butterflies. Our results support the behaviour-at-boundaries hypothesis, with 'softer' boundaries for woodland landscapes. Because the butterflies were reared in a common environment, the observed behavioural differences rely on heritable variation between populations from woodland and agricultural landscapes.     

Multiple causes of sexual segregation in European red deer: enlightenments from varying breeding phenology at high and low latitude

Sexual segregation outside the mating season occurs in most species of sexually dimorphic ungulates and has been extensively described in the literature, but the mechanisms causing segregation are still debated. The detailed pattern of sexual segregation throughout the year has rarely been presented for mammals, and no study, to our knowledge, has used latitudinal-related variation in breeding phenology to shed light on the underlying mechanisms. Recent methodological developments have made it possible to quantify separate components of segregation (social, habitat) and activity synchrony in animal groups, but these major improvements have so far been little used. We observed European red deer year round at two widely different latitudes (France and Norway) and tested three different mechanistic hypotheses of segregation related to: (i) predation risk; (ii) body-size-related forage selection; and (iii) activity budget. Habitat segregation peaked during calving in both populations and dropped rapidly after calving. Females with calves were more segregated from males than were females without calves, pointing to a key role of antipredator behaviour even though large predators are absent in France and extremely rare in Norway. However, at both sites individuals also grouped with their own sex within habitat types (i.e. social segregation), and individuals in mixed-sex groups were less synchronized in activity type than individuals in either unisex male or unisex female groups, suggesting that differences in activity budgets are involved. Social segregation peaked during calving and was lowest during the rut (indicating aggregation) in both populations; these activities occurred one month later in the Northern populations, corresponding well with known differences in breeding phenology. We conclude that latitude-dependent breeding phenology shapes the seasonal pattern of sexual segregation and that sexual segregation in ungulates has multiple causes.     

大兴安岭林区驯鹿夏季喜栖生境的生态特征

2012和2013年7—8月采用样线和样带调查结合的生境调查方法,对分布于我国大兴安岭西北麓的驯鹿的夏季偏好生境开展了对比研究,并对样地的海拔和乔木郁闭度等23个生境变量进行测量及比较.结果表明: 与对照样地相比,夏季驯鹿利用样地的海拔(926.9±0.8 m)、乔木郁闭度(17.9%±2.4%)、乔木胸径(35.5±2.1 cm)、乔木高度(8.2±0.5 m)、乔木密度(6.9±0.5 株·400 m^-2)及树桩数(1.3±0.2 个·400 m^-2)较大,但灌木均高(54.2±2.0 cm)较小(P<0.01).驯鹿夏季偏好选择坡度较缓的中坡生境,因其夏季喜栖生境的水源较多、隐蔽度和避风性较差,距人为干扰距离和牧民点距离均较远.主成分分析(PCA)表明,干扰强度(由距人为干扰距离及距牧民点距离变量组成)、乔木特征(由乔木的高度、密度、胸径和郁闭度变量组成)、地理特征(由坡位、坡向和土壤湿润度变量组成)、食物多度(地表植被盖度和灌木盖度变量构成)、开阔度(由隐蔽度和避风状况变量构成)和坡度是影驯鹿夏季生境选择的重要因素.这6个因子提供了70.7%的累积贡献率,主要反映了驯鹿夏季在喜栖生境选择中的食物资源、抗干扰性和反捕需求.从夏季喜栖生境的生态特征来看,驯鹿并没有被完全驯化.在保护和管理实践中,应使驯鹿种群及其核心栖息地免受高强度的人为干扰.     

Sexual segregation by Masai giraffes at two spatial scales

In this paper alternative explanations for observed patterns of sexual segregation by giraffes are examined at two spatial scales: within-habitats and within-landscape. Habitats are defined as recognizable plant associations and the landscape as the collection of all available habitat types. The study was conducted in Mikumi National Park, Tanzania. At the within-landscape level, all sex and age classes of giraffes exhibited high degrees of preference for riverine habitats. Sex differences in habitat selection were mostly due to females with young, who tended to select open floodplain habitats in which their vigilance time was lowest. Males, and females without young, preferred more heavily-wooded habitat. Habitat preferences were not related to observed habitat-specific forage intake rates for either males or females. Within habitats, male and female giraffes selected different feeding heights, males feeding higher in the canopy than females. Females showed a strong tendency to generalize with respect to feeding height. It is suggested that a sexual dimorphism–body size hypothesis provides a parsimonious explanation for the observed feeding height selection patterns, whereas a reproductive strategy hypothesis can explain sex-differences in habitat selection patterns within the landscape.     

Are hybridogenetic complexes structured by habitat in water frogs?

The success and the evolutionary fate of hybridogenetic lineages are explained by both a generalistic heterosis hypothesis and an alternative hypothesis, the habitat segregation hypothesis. Because such hypotheses have rarely been tested at the level of whole habitats, our aim was to compare performances of two taxa within a hybridogenetic complex across diverse natural habitats. We took advantage of the waterfrog hybridogenetic complex (Rana esculenta and R. lessonae) by rearing tadpoles in natural contrasted habitats by means of enclosure experiments. We also monitored the frequency of each taxon in the waterfrog assemblages that naturally breed in the studied ponds. The hybridogenenetic taxon showed no evidence of broader tolerance as growth, development and physiology strongly varied in response to environmental heterogeneity. Our study reveals a differential success of the hybridogenetic taxon and its sexual host among environments. Moreover, hybridogenetic taxa rarely dominated the sexual species in natural assemblages. Consequently, our results show that the generalistic model does not explain the success of hybridogenetic lineages, but rather support the habitat segregation, among other alternative concepts.     

Additive Partitioning of Diversity Reveals No Scale-dependent Impacts of Large Ungulates on the Structure of Tundra Plant Communities

Large herbivores can change ecosystem functioning by impacting plant diversity. However, although such impacts are expected to be scale-dependent in ecosystems with wide-roaming ungulates, scaling issues rarely enter empirical assessments. We here test the hypothesis that the impact of increased reindeer abundance on plant diversity in alpine tundra is scale-dependent. Based on potentially high productivity of the focal habitat units and hence the possibility of positive grazer impacts on plant diversity we predicted higher α and β diversity at the habitat scale where reindeer densities are high. We also explored whether there were differences in diversity patterns at larger scales, including the scale of reindeer management districts. We estimated grazing disturbance as high versus low reindeer density in selected districts (a total extent of 7421 km²) of Northern Norway where reindeer-induced vegetation shifts are debated. We focus on dominance patterns because they can quantify the vegetation state and thus performed additive partitioning of Simpson diversity on multiple scales assessing also species’ contributions to diversity. Contrary to our predictions, we found only weak scale-dependent effects of reindeer grazing on plant diversity. Under high reindeer densities there was evidence for a landscape-scale homogenization of the vegetation, but the predicted α and β diversity increases at the habitat scale were not found. Consistently through all scales considered, four shrub species contributed the most to plant diversity. These results contradict the idea that reindeer at high stocking densities induce shifts in plant species dominance in productive habitats. We conclude that context-dependencies such as spatial scales of management units and habitat types need to be explicitly considered in evaluations of the impacts of large ungulates on plant diversity.     

Sexual segregation in Tibetan gazelle: a test of the activity budget hypothesis

Tibetan gazelle Procapra picticaudata , a threatened endemic species on the Qinghai-Tibet Plateau, is a sexually dimorphic ungulate. Males are larger than females and both sexes live in different groups, except during the rutting season. In order to test the activity budget hypothesis, which was proposed to explain social segregation in ungulates, we studied the activity budget of male and female Tibetan gazelles during the summers of 2005 and 2006. The activity budget hypothesis predicts that females spend more time feeding and group members synchronize their activities more in the same-sex groups than in the mixed-sex groups. We found that females and males of Tibetan gazelle had different activity budgets; females spent significantly longer time feeding and comparably less time on other activities. Activity synchronization indexes between female groups and male groups were similar (female groups: 0.81±0.17, male groups: 0.80±0.19), whereas both of them were significantly higher than that of mixed-sex groups (0.58±0.29). These results suggest that although sexual segregation might be caused by multiple mechanisms, the activity budget hypothesis about sexual segregation is supported by our studies of Tibetan gazelle on the Qinghai-Tibet Plateau.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

The interaction between bird predation and plant cover in determining habitat occupancy of darkling beetles

Tenebrionid beetles in the Negev Desert exhibit size-related habitat segregation, with larger species found in denser cover. Size-dependent predation by birds has been suggested as the mechanism behind this habitat segregation. Two predictions of this hypothesis were tested: (1) plant cover reduces the predation efficiency of birds upon large tenebrionids, and (2) birds prefer larger species. Both predictions were supported: plant cover reduced predation rate by the most common spring and summer predatory birds: white storks ( Ciconia ciconia ) and stone curlews ( Burhinus oedicnemus ), in cage experiments. Results from preference experiments suggest that tenebrionid species can be divided according to their profitability as prey. Large species are the most profitable, medium-sized species are less profitable but still acceptable and small species are unprofitable and therefore ignored. Field observations demonstrated that the well-vegetated wadi habitats are dominated by large and small species whereas acceptable , medium-sized species are under-represented in this habitat. The results of the cage experiments indicate possible apparent competition between the large profitable and the medium acceptable tenebrionid species in the wadis. Aggregative response of predators in the profitable habitat is suggested as the mechanism leading to truncated distribution of prey species. Large profitable species are refuge-dependent, medium-sized acceptable species use enemy free space and small species are predator independent.