ASSESSMENT OF POTENTIAL BIAS WITH SNOWSHOE HARE FECAL PELLET-PLOT COUNTS

Abstract: The fecal pellet-plot method has been used extensively for snowshoe hare (Lepus americanus) population studies across the species' range, but potential biases associated with the technique have not been addressed adequately. We studied hare pellet-plots in northern Idaho to quantify pellet decomposition rates across environmental gradients, and conducted feeding trials on captive hares to assess the role of diet on pellet production rates. We found that across our study area pellet numbers tended to be higher on plots with high vegetative cover, which likely was a reflection of hare habitat choice rather than lesser pellet decomposition in such habitat. A pellet decomposition experiment indicated that pellet persistence was negatively related to moisture level, and that pellets produced by hares during summer decomposed more quickly than those from winter. We found that only 19% of fecal pellets collected from plots located across northern Idaho were produced by hares during winter. There was a correlation between pellet numbers from plots that were pre-cleared 1 year earlier and estimated numbers of hares on 6 study areas. A similar correlation was lacking for pellet counts from uncleared plots, implying that hare population estimation via pellet-plot counts should involve plot pre-clearing. In captive studies, juvenile hares produced slightly fewer pellets per day per gram of food ingested than adults, but pellet production was similar across diets comprised of 10 different browse species. We conclude that for our study area the fecal pellet-plot method may be subject to notable pellet decomposition bias, and therefore recommend that use of the method elsewhere across the species' range be preceded by assessment of both the pellet-hare density relationship and pellet decomposition rates across habitats.     

A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate

1. European hares Lepus europaeus have declined throughout Europe since the 1960s. Possible reasons for this include agricultural intensification and changes in climate and predator numbers, but no clear consensus has been reached as to the relative importance of each of these. We aimed to identify factors associated with high and low hare numbers throughout Europe, to determine which could have caused population declines. 2. Results of 77 research papers from 12 European countries were summarized. Relationships between hare density and demographics and habitat, climate, hunting and predator variables were examined and quantified where possible. Temporal changes in factors identified as being associated with high or low numbers of hares were then examined to see if they could explain population declines. 3. Data from pastural habitats were limited, but densities of hares were low. Arable habitats had higher densities than mixed areas in spring, unless farming was intensive in which case densities were similar. In autumn the two habitats had similar densities. Field size, temperature, precipitation and hunting had no effect on density throughout Europe. Fecundity was affected by climate. 4. Arable land, various crops, fallow habitat and temperature were positively associated, and monoculture, precipitation and predators negatively associated with hare abundance. The relationship of field size, pasture and woodland with abundance depended on spatial scale. 5. Habitat changes caused by agricultural intensification are the ultimate cause of hare population declines. Effects of changes in climate or predator numbers are magnified by the loss of high-quality year-round forage and cover. Further research is required on how habitat changes affect fecundity and survival, and to identify which parameters have the greatest impact on population numbers. Farmland management policies that target the re-establishment of some of the habitat diversity lost within fields, farms and landscapes will help to reverse the decline of the European hare.     

Relationship Between Fecal Pellet Counts and Snowshoe Hare Density in Western Wyoming

ABSTRACT Snowshoe hares (Lepus americanus) are an important prey species for Canada lynx (Lynx canadensis) and are considered critical for lynx population persistence. Determination of snowshoe hare distribution and abundance is needed by land management agencies for lynx conservation. An accepted approach for estimating snowshoe hare abundance is the use of fecal-pellet plot counts. Locally derived regression equations are preferred for accurate calibration of pellet counts to snowshoe hare density due to local differences in pellet deposition and decomposition. We used linear regression to examine correlations between snowshoe hare density, as determined by mark–recapture estimates, and pellet plot counts on both uncleared plots and annually cleared plots on the Bridger-Teton National Forest, western Wyoming, USA. We found significant correlations between snowshoe hare density estimates and fecal pellet counts for both uncleared and annually cleared pellet counts; however, the relationship was stronger (higher r) when using pellet counts from annually cleared plots. In addition, we found that adjusting the buffer size by omitting hard habitat edges (not used by hares) around trapping grids improved correlations between snowshoe hare density and fecal pellet counts for both uncleared plots and annually cleared plots. Though precision is sacrificed when using uncleared plots, they may be useful as a coarse index of habitat use by snowshoe hares. Our derived regression equations may be useful to identify important foraging habitat for Canada lynx in western Wyoming. Land managers responsible for conserving snowshoe hare habitat in western Wyoming may use these equations to monitor changes in hare populations among habitats and during prescribed management actions.     

Spring and autumn habitat preferences of active European hares (Lepus europaeus) in an agricultural area with low hare density

The number of European brown hares (Lepus europaeus) has been declining throughout much of Europe since the 1960s. Consequently, many studies have focused on analysing habitat selection of European hares in order to improve the suitability of the habitat for this species. Habitat preferences of European hares are known to be affected by hare density, but most studies have been conducted in agricultural areas where hare densities were medium to high. Finding habitat preferences at high densities is difficult as most available habitats are occupied. In addition, in agricultural areas, field size might influence the hares’ habitat selection because it affects the distribution and availability of certain habitat types. However, most studies relate to areas with large field sizes. In this study, we analysed the habitat preferences of European hares in spring and autumn during the activity period, in the early hours of the night, in an agricultural area with low hare density and small average field size using Chesson’s electivity index. Moreover, we focused on the question whether two different habitat classifications varying in their specificity might cause contradictory results regarding European hares’ habitat preferences. Our results show that in this agricultural area with low hare density, European hares avoided several habitat types which were preferred in other study areas with higher hare densities. Therefore, we assume that hare density has an influence on the species’ habitat selection. In contrast, the small average field size of our study area seemed not to have an effect on hare habitat preference. Furthermore, by pooling habitat types into broader groups, substantial information was lost in some categories. Hence, for some categories, e.g. grassland or agricultural crop land, more detail might be needed than for others, such as urban areas, when analysing hares’ habitat selection. In conclusion, our results imply that studies on habitat preferences have to be conducted in areas with low hare density to be able to gain knowledge on the species’ habitat requirement and hereinafter improve the suitability of the habitat for this species.     

The diet of the mountain hare (Lepus timidus hibernicus) on coastal grassland

Across most of their range in Europe, mountain hares are usually restricted to upland areas with poor food quality. In these areas they generally feed on browse species such as heather or twigs and barks of trees. On lowland areas in Europe, with better food quality, the mountain hare is replaced by the brown hare ( Lepus europaeus ) which feeds predominantly on greasses. This khas led some authors to conclude that mountain hares are primarily adapted for browsing. In the absence of brown hares in Ireland, mountain hares are found on a wide variety of habitats including grassland. On grassland, their diet consists almost exclusively of grasses, up to 94% of their annual diet, which is more than has been reported for brown hares on similar habitat. Based on this evidence, and other work, it is proposed that the mountain hare in primarily a grazing animal and competitive exclusion by brown hares may underlie much of their present distribution in Europe.     

Home range use by the European hare (Lepus europaeus) in a structurally diverse agricultural landscape analysed at a fine temporal scale

An animal’s home range use is influenced by the landscape type. European hare (Lepus europaeus) home ranging behaviour has been studied only in agricultural areas with medium to large fields. In agricultural areas with small fields, European hares’ locomotor behaviour is expected to be more localised. We tracked nine European hares by means of global positioning system (GPS) and very high-frequency (VHF) collars during summer in an agricultural area with small fields in Lower Austria. In particular, we analysed the hares’ space use at a fine temporal scale, such as when they were active and resting within single 24-h periods. Furthermore, we compared data (day–day distances and day–night distances travelled) calculated from GPS and VHF telemetry. Home ranges were smaller, and the distances between areas used for activity and inactivity were shorter, in this agricultural area with small fields than has ever been measured in other agricultural areas with larger fields. Both active and inactive European hares expressed a preference for areas near field edges. Our findings suggest that with GPS, it is possible to distinguish between the movement path and the relative location of distinctly used areas within an animal’s home range, whereas with VHF these two parameters may be difficult to separate. In conclusion, our results show that in areas where resources are easily accessible, such as in agricultural areas with small fields, the European hare is able to reduce its home range size to almost half of the minimum size that has been recorded so far in other habitats. As small home ranges involve less energy expenditure for movement, our results suggest that animals living in agro-ecosystems may benefit from small fields.     

Limited effect of hare grazing and short-term climatic variations on the most common alpine vegetation community in the Snowy Mountains,Australia

Background: In the alpine zone of the Snowy Mountains, grazing by mammals is limited. However, introduced European hare numbers have increased since the 1970s.

Aims: To estimate the density of hares and hence grazing pressure among years. To assess the response of biomass, vegetation height and composition to a cessation of hare grazing.

Methods: We used indices of hare abundance based on spotlighting and counts of hare pellets on a transect. The effect of hare grazing on tall alpine herbfield was assessed by using 15 paired exclosure and control quadrats for six years.

Results: The indices of hare abundance suggested densities similar to those in upland areas of Britain. Grazing did not affect the composition, cover of herbs or graminoids or, for 2010, vegetation height or biomass. Variation in vegetation and hare numbers among years was not correlated with climatic variables. Observations of selective grazing suggested that impacts on vegetation may be localised and restricted to certain species. Prior analyses of hare pellets indicated that hares might spread seed of native and exotic species.

Conclusions: Hares are having no general effect on tall alpine herbfield but may affect certain plant species via selective grazing or by spread of viable seed.     

The use of block counts,mark-resight and distance sampling to estimate population size of a mountain-dwelling ungulate

Population size estimates represent indispensable tools for many research programs and for conservation or management issues. Mountain ungulates in open areas are often surveyed through ground counts that normally underestimate population size. While the use of sample counts is desirable, few studies have compared different probabilistic approaches to estimate population size in this taxon. We compared the size estimates of a male population of Alpine chamois using mark-resight and line transect sampling methods, while block counts were used to obtain the minimum number of males alive in the study area. Surveys were conducted within the Gran Paradiso National Park (Italy), in August–September 2013, using block counts along purposely selected trails and vantage points, mark-resight over 5 consecutive resightings from vantage points and trails, and line transect sampling along 12 transects repeated 8 times. Block counts yielded a minimum number of males alive in the population of n = 72 individuals. This value was greater than the upper bound of the 95 % confidence interval achieved using line transect sampling {n = 54, CV = 14 % [95 % CI (40, 71)]} while mark-resight yielded a more realistic result of n = 93 individuals {CV = 18 % [95 % CI (63, 137)]}. Our results suggest that line transect sampling performed poorly in the Alpine environment, leading to underestimates of population size, likely due to violations of some assumptions imposed by the rugged nature of the terrain. The mark-resight yielded lower precision, possibly due to the limited number of marked individuals and resighting occasions, but it provided robustness and accurate estimates as marks were evenly distributed among animals.     

Highest densities of mountain hares (Lepus timidus) associated with ecologically restored bog but not grouse moorland management

Over the last 20 years, ecological restoration of degraded habitats has become common in conservation practice. Mountain hares (Lepus timidus scoticus) were surveyed during 2017–2021 using 830 km of line transects in the Peak District National Park, England. Historically degraded bog areas were previously reported having low hare numbers. Following bog restoration, we found hare densities of 32.6 individuals km⁻², notably higher than neighboring degraded (unrestored) bog with 24.4 hares km⁻². Hare density on restored peatland was 2.7 times higher than on bogs managed for grouse shooting at 12.2 hares km⁻² and 3.3 times higher than on heather moorland managed for grouse shooting at 10.0 hares km⁻². Yearly estimates varied most on habitats managed for grouse, perhaps indicative of the impact of habitat management, for example, heather burning and/or possible hare culling to control potential tick‐borne louping ill virus in gamebirds. Acid grassland used for sheep farming had a similar density to grouse moorland at 11.8 hares km⁻². Unmanaged dwarf shrub heath had the lowest density at 4.8 hares km⁻². Hare populations are characterized by significant yearly fluctuations, those in the study area increasing by 60% between 2017 and 2018 before declining by ca. 15% by 2020 and remaining stable to 2021. During an earlier survey in 2002, total abundance throughout the Peak District National Park was estimated at 3361 (95% CI: 2431–4612) hares. The present study estimated 3562 (2291–5624) hares suggesting a stable population over the last two decades despite fluctuations likely influenced by weather and anthropogenic factors. Mountain hares in the Peak District favored bog habitats and were associated with restored peatland habitat. Wildlife management should be cognizant of hare density variation between habitats, which may have implications for local extinction risk.     

NIGHT COUNTS OF HARES AND OTHER ANIMALS IN EAST AFRICA

Night counts of hares and other animals were made on a disused airstrip in the Queen Elizabeth National Park, Uganda and on the road between Magadi and Nairobi in Kenya. Lepus crawshayi is the only species of hare which occurs in the Queen Elizabeth Park but L. capensis is also found in Kenya. No regular seasonal fluctuations were found in the number of hares recorded due probably to the continuous breeding of these equatorial species. The number of waterbuck seen in Uganda on moonlit nights was significantly greater than that counted on dark nights. This is not thought to be due to better viewing conditions under a full moon since the phenomenon was not found with other species. The number of hares recorded varied inversely with the number of other mammals present but bore no relationship with the number of birds. It is concluded that night counts can be useful in revealing population trends but not for assessing total numbers or for comparing densities in different areas.     

The recent expansion of the brown hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) in Sweden with possible implications to the mountain hare (<Emphasis Type="Italic">L. timidus</Emphasis>)

The brown hare (Lepus europaeus) expanded its Swedish distribution since the 1980s northwards and locally to new areas within its former range. Of 115 brown hare populations within the former range reported in a hunter enquiry, those established after 1980 were situated higher above the sea level than older ones and higher than neighbouring (<50 km) older populations. Reports on increased use of forest habitats by brown hares were equally frequent among recent and older populations, suggesting a process promoted solely by less harsh winters. Supposed hare hybrids were more often reported from hunting grounds with recent brown hare establishment, i.e. where the species expands in time and in space. In a 27-year dataset on brown hare observations, the recent increased use of forest habitats was supported in that maximum distances to agricultural land for brown hare sightings were higher in mild winters, whereas the proportions of the annual observations made during winter were lower. In 40-year bag records from two Swedish counties, the dynamics of the mountain hare (Lepus timidus) responded positively to snow parameters, whereas brown hares responded negatively. We suggest that the state of mountain hare populations primarily depends on winter conditions and predation pressure, whereas possible effects of hybridization are unclear. If winter conditions remain as in the last 15 years, mountain hare numbers are not likely to increase in southern Sweden, whereas the brown hare may expand even further. In either case, hybrids will occur in sympatric areas in frequencies probably related to the density of the respective true species.     

Temporal dynamics of European brown hare syndrome infection in Northern Italian brown hares (Lepus europaeus)

The progressive decline in the hare population across Europe has been associated with the occurrence of European brown hare syndrome (EBHS), a highly contagious disease considered endemic in all European countries. This study aimed to evaluate the in-field temporal dynamics of European brown hare syndrome virus (EBHSV) infection in wild European brown hares (Lepus europaeus) and to test the influence of population density on EBHS seroprevalence. A total of 512 blood samples were collected from free ranging hares captured for restocking in seven different areas of the province of Brescia (Northern Italy) during seven consecutive years (2006–2013) and tested using a competitive ELISA. A generalized linear mixed model estimated the yearly effects of population density on EBHS prevalence. Of the 512 tested, 344 (67.2 %) tested positive for EBHSV antibodies, with the annual seroprevalence ranging from 94.3 to 35.8 %. The prevalence was 3.303 times higher in areas with a density of over 15 hares/km² and declined over the years. The results indicate the ongoing transmission of the virus in the tested brown hare population. Since the eradication of EBHS in a wild population is not feasible, a strategy aimed at promoting the endemic stability of the virus through density-dependent mechanisms could be applied; however, this seems more difficult in practice than in theory and would most likely require a very high density of brown hares.     

Methods of studying the distribution, diversity and abundance of birds in East Africa—some quantitative approaches

In this paper we compare the use of transect counts with a simpler method of investigating bird diversity and numbers, particularly in terrestrial habitats, both natural and non‐natural. Transect Counts (TCs) have long been widely used, whereas Timed Species Counts (TSCs), which estimate relative abundance, are comparatively untried. We find that TSCs give results which are comparable to those from TCs in most respects, except that they can only be used indirectly for estimating population densities, and they give different measures of diversity. However, TSCs generate data on many more species much faster than do TCs and are therefore more cost‐effective in most situations. In particular, TSCs are useful for community studies. We show, for example, that in natural habitats bird populations are positively correlated with the amount of woody vegetation, but not with rainfall. Diversity too increases with woody vegetation. Because TSCs are simple, more of them can be made for a given input of time, and hence more distributional data are obtained as an additional benefit.     

山西省草兔的一些生态资料

山西省草兔的一些生态资料卢欣，申守义，高尚文（山西省生物研究所太原０３０００６）关键词草兔，粪便样方，益害分析，天敌和寄生虫草兔（Ｌｅｐｕｓｃａｐｅｎｓｉｓ）是我国最重要的小型狩猎动物之一，但其生态仅有零星报道［１，２］。１９８７－１９９２年，我们在...     

The effect of landscape structure on the distribution of brown hare Lepus europaeus in farmlands of Germany and Poland

Habitat management should be an important part of the brown hare (Lepus europaeus) conservation, but the habitat requirements of this species are not fully recognised. The aim of our research was to estimate these requirements by analysing the effect of various agricultural landscape structure features on the distribution of hares in five agricultural areas in Germany and Poland. The local density of hares was assessed in the spring and autumn of 2006 by using the method of spotlight–strip counts on 9–15 subareas in each research region. The structure of agricultural landscape has been described for each subarea: the share of grain, other crops and grasses as well as the density of crop edges and uncultivated places with wild vegetation. The density of hares was considerably higher in Germany than in Poland (18.8–48.4 vs. 4.1–9.5 indiv./km²). The hare density was positively correlated with non-grain crops in an area, with crop edges in two areas and with wild vegetation without trees in two areas, and negatively correlated with grassfields in two areas. The occurrence of wild vegetation without trees affected the hare density only in the study areas, where this habitat was relatively rare (<3 km/km²). It was suggested that proper projects aimed at habitat management for brown hares should be elastic, i.e. the projects should be modified depending on the structure of local landscapes. Moreover, the protection and creation of structures with wild vegetation among cropland seem to be considerable methods of brown hare or generally wildlife conservation; therefore, such measures should be an important part of agro-environmental packages.     

Sinks without borders: snowshoe hare dynamics in a complex landscape

A full understanding of population dynamics of wide-ranging animals should account for the effects that movement and habitat use have on individual contributions to population growth or decline. Quantifying the per-capita, habitat-specific contribution to population growth can clarify the value of different patch types, and help to differentiate population sources from population sinks. Snowshoe hares, Lepus americanus , routinely use various habitat types in the landscapes they inhabit in the contiguous US, where managing forests for high snowshoe hare density is a priority for conservation of Canada lynx, Lynx canadensis . We estimated density and demographic rates via mark–recapture live trapping and radio-telemetry within four forest stand structure (FSS) types at three study areas within heterogeneous managed forests in western Montana. We found support for known fate survival models with time-varying individual covariates representing the proportion of locations in each of the FSS types, with survival rates decreasing as use of open young and open mature FSS types increased. The per-capita contribution to overall population growth increased with use of the dense mature or dense young FSS types and decreased with use of the open young or open mature FSS types, and relatively high levels of immigration appear to be necessary to sustain hares in the open FSS types. Our results support a conceptual model for snowshoe hares in the southern range in which sink habitats (open areas) prevent the buildup of high hare densities. More broadly, we use this system to develop a novel approach to quantify demographic sources and sinks for animals making routine movements through complex fragmented landscapes.     

How accurate are single site transect data for monitoring butterfly trends? Spatial and temporal issues identified in monitoring Lasiommata megera

Multiple transect counts following Butterfly Monitoring Scheme (UKBMS) guidelines and Jolly–Seber estimates of population size were used to monitor the abundance of second generation Lasiommata megera on a single site in southern England. The two methods resulted in different patterns of emergence being detected. The proportion of the population (estimated by Jolly–Seber) recorded with transect counts depended on the time of day and weather with afternoon transect counts best recording the trend in abundance over the flight period, but even then counts recorded a variable fraction of the population (6.2–51.3%). Increasing the frequency with which transect counts are carried out per week reduced variation and increased the fit of transect counts to Jolly–Seber generated population estimates. However, indices of abundance generated from randomly selected transect counts for L. megera within sampling weeks varied 4-fold and indices for other butterfly species were also highly variable. For L. megera, transect count variability is attributed to non-representative placement of the transect route and changes in the behaviour and spatial distribution in relation to population size and season. We suggest that transect counts need to be fully validated before the data are used to monitor changes of butterfly populations at individual sites.     

Population density,roads and altitude influences on spatial distribution of hares positive to EBHSV

Understanding of the ecology of infected animals facilitates disease risk assessment and is also crucial for wildlife conservation. Relatively little is known about the spatial distribution of infected wild mammals in relation to environmental factors. In neighboring Mediterranean ecosystems 250 European brown hares (Lepus europaeus) were collected and examined with RT-PCR to detect European Brown Hare Syndrome Virus (EBHSV). Multivariate statistics and Geographical Information System (GIS) analysis were applied to estimate spatial patterns of biotic and abiotic factors and human activities as determinants of EBHSV positivity. Hare population abundance was estimated using faeces counts and belt drive censuses. The study showed that EBHSV infected hares had widespread distribution even in isolated areas. However, EBHSV infection prevalence was higher in areas with higher hare abundance, closer to paved road networks and at lower altitudes. The risk map revealed the potential distribution of EBHSV-infected hares. This study shows that host abundance and landscape influence the ecology of the disease, a finding that should be taken into account in future studies. The management of harvest and restocking of hares is also discussed for population conservation.     

Sampling tiger ungulate prey by the distance method: lessons learned in Bardia National Park, Nepal

Because tiger Panthera tigris numbers are regulated by their prey base, prey abundance needs to monitored and estimated reliably. Recently, distance sampling has been adopted as the most appropriate method and is now becoming the standard monitoring protocol in all tiger range countries in south Asia. However, the accuracy of the density estimates generated by this method has not been assessed. From total counts within habitat blocks, we obtained accurate density estimates of ungulates within three main habitats in Bardia National Park, Nepal. We then applied the distance sampling method in the same habitats and compared the results. Distance sampling on foot in dense habitats (riverine forest and tallgrass floodplain) violated method assumptions, and sampling from vehicle along roads gave biased estimates. Sampling from elephant back worked well in all habitat types, but owing to their behaviour, the density of barking deer Muntiacus muntjak was underestimated. The accuracy of the estimates varied with sampling effort; for the very abundant chital deer Axis axis , estimates varied markedly at <200 animal observations, but converged at larger sample sizes to a similar point estimate as intensive block counts when approaching 300 observations. For the less abundant species, with <20 observations along >100 km of transect lines, the confidence intervals were quite high, and, hence, of limited value for detecting short-term populations trends. It is therefore difficult to obtain accurate density estimates of rare species by the distance method. In areas consisting of dense habitats, we recommend that the food base of tiger be estimated by distance sampling from elephant back, not on foot, directed at the main and most abundant prey species. For rare species, encounter rates obtained simultaneously may then serve as indices of relative abundances.     

Implications of shared predation for space use in two sympatric leporids

Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.