Diet of the mara (Dolichotis patagonum), food availability and effects of an extended drought in Northern Patagonia (Mendoza,Argentina)

The mara is a near threatened Caviomorph, endemic to Argentina. Studies on feeding ecology allow assessment of the dietary adaptability of maras to habitat changes. The mara diet and food availability on two sampling sites, belonging to distinct landscapes of Northern Patagonia, were estimated using microhistological analysis and point-quadrat transects, over four seasons, and besides during an extended drought. Significant differences were detected by Kruskall-Wallis ANOVA, feeding selection by the χ² test, and dietary preferences by Bailey's confidence interval. Grasses dominated food availability, with Panicum and Poa as major species, followed by shrubs and scarce forbs. Plant cover and forbs increased in spring and summer. The drought caused a strong decrease in plant cover and proportion of grasses. Maras ate all grass species, most forbs and several shrubs. Grasses dominated the diet, with Poa and Panicum being the major species, supplemented by the shrubs Lycium and Prosopis. Maras ate more grasses and forbs in spring and summer, and shrubs in autumn and winter. More shrubs and forbs, and less grasses, were eaten during the drought. Plant categories were used selectively only in autumn and winter, and in the drought period, with preference for shrubs and avoidance of grasses. Bromus, Poa, Plantago and Prosopis were preferred, and Panicum avoided. The mara qualified as a grazer but shifted to a mixed feeder during the drought. Dolichotis patagonum shared habitats with several big and medium-sized herbivores and showed the highest dietary similarities with plain vizcachas, brown hares and horses. Protective measures for natural habitats are needed, given that increasing impacts on food resources and habitat quality could be threats to the survival of maras and other wild vertebrates in Northern Patagonia.     

Interactions between hare and brent goose in a salt marsh system; evidence for food competition?

In this study we accumulate evidence that brown hare competes with brent goose for food resources in a temperate salt marsh. We show that both species overlap in habitat use and share food plants. The two herbivores mainly used the common habitat at different times of the day, with hares active in the dark and geese during the daylight. During the morning and evening, however, the habitat was exploited simultaneously. Food availability was manipulated by excluding brent geese on both small-scale (30 m²) and large-scale (0.96 ha) plots, while hares had free access everywhere. Exclusion of brent geese enhanced the level of utilisation by hares in both Festuca and Puccinellia dominated marshes, which are among the most intensively grazed parts of the salt marsh. The increase in hare grazing pressure following goose exclusion was stronger, when the adjacent control plots had attracted more goose visitation. When geese were excluded, the decrease in Festuca consumption by geese was completely matched by increased hare grazing, while for Puccinellia only part of the `surplus' was harvested. Enhanced levels of hare utilisation were not due to geese interfering directly with hare, nor due to hares avoiding goose droppings. Considering the interaction from the other perspective, hares were observed to disturb geese effectively in every spring. This might have reduced exploitation by geese of the shared resources. On the basis of our experimental results, we conclude that in this salt- marsh system competition for food with brent geese plays a role in the habitat use of hares, and that hares can reduce goose exploitation of shared habitats. Received: 30 March 1998 / Accepted: 6 July 1998     

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.     

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.     

Molecular Approaches Revealing Prehistoric, Historic, or Recent Translocations and Introductions of Hares (Genus Lepus) by Humans

Although only of medium size, and thus of little nutritional value compared to big game such as mammoths and ungulates, hares (Lepus spp.) probably have always been a food source for humans, as documented in archaeological finds. Nowadays, hares, particularly such species as the brown hare (L. europaeus), are among the most important game species in many European countries. For hunting, perhaps religious reasons, and in connection with certain myths, hares have been and are still being intentionally translocated. Ancient translocations by humans can be inferred from the presence of hares on islands that had no mainland connections, at least during the Pleistocene, the major evolutionary period of the genus Lepus. We review some of the literature on anthropogenic translocations of hares. We focus on three examples [the brown hare (L. europaeus), the Corsican hare (L. corsicanus), and the Sardinian hare (L. capensis)], where some molecular data could be used to trace the translocation routes and possible origins of introduced hare populations. Certain molecular marker systems, such as sequences of the hypervariable part I (HV-1) of the mitochondrial control region, show high variability in hare species and are thus promising for tracing both recent and ancient origins of translocated hares. Some other molecular marker systems as well as caveats connected with the use of such marker systems in the genus Lepus are also discussed.     

Foraging behavior and spatial use of a rock specialist: the southern vizcacha (Lagidium viscacia), and the exotic European hare (Lepus europaeus) in rocky outcrops of northwestern Patagonia, Argentina

The southern vizcacha (Lagidium viscacia) and the exotic European hare (Lepus europaeus) are two medium-sized herbivores that inhabit rocky outcrops in Patagonian steppe. These species overlap in diet and spatial use at medium distances from rocky outcrops in summer. We evaluated the spatial use through feces distribution in winter and determined seasonal foraging intensity in relation to the distances from rocky outcrops in order to elucidate how these herbivores use food and spatial resources in food scarcity periods. The vizcacha utilized the habitat close to rocky outcrops (<40 m) independent of season, while the hare exploited the space more widely, especially distances >40 m. However, in winter, at medium distances from rocky outcrops, there was partial spatial overlap because hares' activities were closer to rocky outcrops. Foraging intensity increased significantly in areas used by the vizcacha closer to rocky outcrops when food availability decreased, and the grasses Stipa speciosa, Poa sp., and Festuca pallescens were strongly foraged. In contrast, foraging intensity showed no changes in further distances to rocky outcrops and more use by the hare. The spatial and feeding behavior of the vizcacha, restricted to vicinity of rocky outcrops, showed high vulnerability to food availability changes. In resource scarcity situations, the spatial opportunistic behavior of the hare and the overlap in diet with the vizcacha constitutes a threat to this native herbivore. It is necessary to monitor populations of hare, since high densities could lead to food competition, impacting the small colonies of the southern vizcacha.     

Hare and vole browsing preferences during winter

The aim with this study was to, under controlled conditions, determine the food preference of mountain haresLepus timidus Linnaeus, 1758 and bank volesClethrionomys glareolus (Schreber, 1780) for a substantial part of the woody plants potentially available for these herbivores during winter. In addition, we compared hare and vole preference patterns. Thirteen woody plant species were simultaneously presented to 9 captive voles and 9 captive hares in preference tests during winter. Consumption by hares from 50 g bundles (one per species) was measured after 3 h, whereas shoot consumption by voles was measured after 12 h. Both hares and voles preferred deciduous species to conifers,Populus tremula, andVaccinium myrtillus being the most preferred species. However, there was considerable variation in palatability among deciduous plant species and only a marginally significant correlation was found between hare and vole preference. One striking differences between hares and voles was that Sorbusaucuparia was the most utilised by voles but the least preferred by hares. In conclusion, deciduous plant species were generally considerably more palatable to hares and voles than conifers, which is consistent with current theories. Nevertheless, the high variation in palatability among deciduous trees and the difference in preference between hares and voles indicate more complex and species-specific patterns with regard to plant— animal interactions. The latter also suggests that hares and voles differ in their dietary adaptations and have different dietary constrains.     

Diet selection of the southern vizcacha (<Emphasis Type="Italic">Lagidium viscacia</Emphasis>): a rock specialist in north western Patagonian steppe,Argentina

The southern vizcacha (Lagidium viscacia) is a rock specialist that inhabits small colonies in isolated rocky outcrops of northwestern Patagonia. This study analyzes its diet selection in relation to food availability, establishes the degree of dietary specialization, and discusses the potential competition with exotic herbivores. Diet composition and food availability were determined in summer and winter in eight rocky outcrops by microhistological analysis of fecal pellets, and food availability was estimated by the Braun Blanquet cover abundance scale. Vegetation cover differences were detected by using a random analysis of variance (ANOVA) factorial block design, and dietary preferences were determined by the confidence interval of Bonferroni. The southern vizcacha showed a specialized feeding behavior despite the consumption of a wide variety of items. Their diet was concentrated on a few types of food, mainly grasses, and the trophic niche was narrow and without seasonal variations. In winter, when food was scarce and of lower quality than summer, diet was dominated by Stipa speciosa, suggesting a selection according to the selective quality hypothesis. Our results (narrow trophic niche, restricted activity near rocky outcrops, and a diet with high proportions of low-quality grasses) showed that the vizcacha is an obligatory dietary specialist, and these characteristics made it highly vulnerable to changes in food availability. In this scenario, overgrazing caused by alien species with similar diets, as the European hare and livestock, could negatively affect their colonies.     

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.     

The distribution of mountain hares Lepus timidus in Europe: a challenge from brown hares L. europaeus?

1. Throughout the most recent glacial period (Weichsel), the mountain hare Lepus timidus had a continuous distribution in the tundra habitat south of the ice‐rim. When the ice retreated, mountain hares colonized deglaciated land, and spread over northern Europe. 2. Since the Weichsel, the mountain hare's distribution in Europe has been gradually reduced and at present comprises Ireland and the Scottish Highlands, high altitudes in the Alps, isolated forests in eastern Poland, most of Fennoscandia and from the Baltic countries eastwards through Russia. Declines during the last century have been observed in Sweden and Russia. 3. This review defines and evaluates causes for this gradual reduction and fragmentation of the mountain hare's distribution, with special focus on interactions with brown hares Lepus europaeus. The relative importance of diseases, predation, cultivation and interactions with other herbivores than brown hares are discussed. 4. A plausible cause of the possible permanent disappearance of mountain hares in Europe appears to be exclusion by interspecific competition and hybridization with, and/or epidemic diseases mediated by, the congeneric brown hare.     

Biochemical genetic variability in brown hares (Lepus europaeus) from Greece

Allozyme variability of 91 brown hares (Lepus europaeus) from seven regions in Greece was compared to existing data of Bulgarian populations to test the hypothesis of the occurrence of specific alleles in Greece, likely stemming from an isolated Late Pleistocene refugial population in the southern Balkans. This hypothesis is particularly suggested by some subfossil Late Pleistocene hare remains in Greece and the reported high mtDNA diversity in Greek hares. Allozymic diversity could be higher in Greek hares than in hares from neighboring regions as a result of the accumulation of variants in a long-lasting Pleistocene refugium. Conversely, Greek hares could exhibit reduced genetic diversity because of long-lasting low effective population sizes during the Late Glacial Maximum and a lower chance of postglacial gene flow from other populations into this rather marginal part in the southern Balkans. Horizontal starch gel electrophoresis of proteins from 35~loci revealed three alleles (Es-1 ^–162, Pep-2 ¹¹⁴, Mpi ⁸⁸) at low frequencies, which were not found in Bulgarian or any other brown hare population. In contrast, some alleles from the populations from Bulgaria and other regions of Europe were absent in the Greek samples. Population genetic statistics indicated only a slight tendency of increased gene pool diversity in Greek hares, little substructuring in Greek and Bulgarian populations, respectively, as well as an only slightly lower level of gene flow between the two neighboring regions, as compared to the gene flow within each region. The results conform to the hypothesis of a Late Pleistocene refugial population in the southern Balkans, with some few specific nuclear gene pool characteristics, but little effect on the overall genetic differentiation between Greek and Bulgarian hares.     

UV-B-induced plant stress as a possible cause of ten-year hare cycles

Predation has been assumed to be a necessary factor in the ten-year population cycle of the snowshoe hare (Lepus americanus) and Canadian lynx (Lynx canadensis). The UV-B-induced plant stress hypothesis, in contrast, predicts that hare performance, especially reproduction, is negatively related to sunspot numbers, because production of UV-B-protective phenolics in food plants in periods of low sunspot activity, when the ozone layer is thin, increases the availability of amino acids and reduces the amount of phenolics that protect against herbivores. In accordance with the UV-B-induced plant stress hypothesis, and despite the absence of predators that have been assumed to be necessary for hare cycles, mountain hare (Lepus timidus) populations in Norway fluctuate in close synchrony with snowshoe hare populations in Alberta and the Yukon, Canada. When adjusting for the phase of the hare cycle, the natality of snowshoe hare in Alberta 1962–1976 was negatively related to sunspot numbers with a time lag of two years. It is concluded that delayed responses to UV-B-induced changes in plant chemistry during the sunspot cycle is a possible cause of ten-year cycles of hares and other herbivores, for example grouse and forest moths.     

Food selection by the guanaco (Lama guanicoe) along an altitudinal gradient in the Southern Andean Precordillera (Argentina)

Wild ungulates like the guanaco are exposed to important changes in climate and plant diversity along altitudinal gradients in the Andes Mountains, such as in the Southern Andean Precordillera where three phytogeographic provinces are present in altitudinal belts. The guanaco’s diet and food availability were seasonally analyzed using microhistological analysis and point-quadrat transects at six sampling sites, representative of the phytogeographic belts along the altitudinal gradient. Plant cover and diversity decreased with growing altitude. Richness of plant species was poorer at the summit than in the lower altitudes, whereas the proportion of species eaten by guanacos increased with altitude. The diet included 77 species. Grasses were preferred and shrubs were avoided all year round. The grass Poa spp. occupied more than 50 % of the diet at all altitudes. Grasses were the main dietary item even at low altitudes, where shrubs constituted the main food available. Decreasing generalism with descending phytogeographic belts agrees with the prediction for altitudinal gradients. The increase of diversity in the diet during the winter decline of plant cover at high and middle altitudes follows that expected from the optimal foraging theory. The winter decline of vegetation and the dietary shift from grazing to browsing proved to be stronger as altitude increases and the climate become more rigorous. Plant species richness, food scarcity, and climate severity are relevant variables to explain altitudinal and seasonal changes in the diet of adaptive ungulates in mountain environments, such as the guanaco in the Southern Andean Precordillera.     

Space use of a non-native species,the European hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis>), in habitats of the southern vizcacha (<Emphasis Type="Italic">Lagidium viscacia</Emphasis>) in Northwestern Patagonia,Argentina

Two medium-sized herbivores with high trophic overlap coexist on rocky outcrops in the Patagonian landscape: the southern vizcacha (Lagidium viscacia), which is a native rock specialist, and the European hare (Lepus europaeus), which is a non-native species. We determined the patterns of space use related to distance from outcrops and analyzed spatial overlap between the two species. There were significant differences between the two species in the use of space adjacent to outcrops. The southern vizcacha mainly uses short and medium distances from the outcrop (up to 40 m), whereas the hare’s greatest activity was recorded at distances greater than 50 m. However, there is a partial overlap at medium distances (30–40 m) among both herbivores. Although, in general terms, there is no significant spatial overlap between hares and southern vizcachas, their biological characteristics and the high dietary overlap between the species allow us to predict that, if resources become scarce, the hare could extend its area of activity, as what happens elsewhere, and exploit food resources near outcrops, increasing the vulnerability of vizcacha colonies.     

Trophic interactions between two medium-sized mammals: the case of the native Dolichotis patagonum and the exotic Lepus europaeus in a hyper-arid ecosystem

As medium-sized herbivores, the exotic Lepus europaeus (European hare) and the native Dolichotis patagonum (mara) have been considered ecological equivalents. These species coexist in Ischigualasto Provincial Park, a hyper-arid ecosystem with scarce food resources. Our objective was to evaluate diet composition, relationship between diets and food availability, and trophic relationships between both herbivores. Collection of feces and vegetation sampling were made in the Mesquite woodland community. Diet composition was analyzed by microhistological analysis of feces. In both seasons, shrub species represented the most abundant cover type in the area, and annual forbs and grasses appeared in the wet season. Herbivores showed similar dietary ecology: shrubs were the main food items along the year, showing a higher plasticity compared to their diets in other ecosystems, where they selected mostly grasses. The mara selected shrubs such as Atriplex sp. and Prosopis torquata, whereas the European hare selected Cyclolepis genistoides, Atriplex sp., and Bulnesia retama. During the wet season, both herbivores supplemented their diets with grasses and annual forbs. In the dry season, there was increased consumption of cacti, such as Tephrocactus sp. The mara and the European hare are likely close ecological equivalents, in terms of dietary similarity, and they showed strong dietary overlap across the dry season (over 60 %). Thus, we can assume the existence of a potential trophic competition between mara and European hare, especially during the season when food resources are scarce. These results can be important for the management of drylands in South America, where populations of threatened herbivorous species, such as the mara, coexist with exotic animals, sharing spatial and trophic resources even in protected areas.     

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.     

Diet selection by hares (Lepus europaeus) in arable land and its implications for habitat management

Populations of European hares (Lepus europaeus) have experienced a dramatic decline throughout Europe in recent decades. European hares are assumed to prefer weeds over arable crops, and weed abundance was reduced by the intensification of agriculture. Therefore, modern agriculture has been blamed as a major factor affecting European hare populations. However, it is questionable whether European hares select weeds at all, as previous studies had major methodological limitations. By comparing availability and use of plants with Chesson’s Electivity Index, we investigated whether the European hare actually feeds selectively on different plants in arable land. Food availability and use were dominated by cultivated crops (e.g. winter wheat, spring barley and sugar beet). Diet selection analysis revealed that in autumn and winter, European hares predominantly preferred cultivated crops (winter wheat) and food items provided by hunters (tubers of sugar beet and carrot). In spring and summer, apart from soy, only weeds (e.g. clover and corn poppy) were positively selected, especially after cereal crops were harvested. We suggest that the decline in European hare populations throughout Europe was facilitated by the decrease in weed abundance. Wildlife-friendly set-asides in arable land have the potential to reconcile the European Union’s Common Agricultural Policy with wildlife conservation.     

Factors related to the occurrence of hybrids between brown hares Lepus europaeus and mountain hares L. timidus in Sweden

Hybridization occurs among many species, and may have implications for conservation as well as for evolution. Interspecific gene flow between brown hares Lepus europaeus and mountain hares L. timidus has been documented in Sweden and in continental Europe, and has probably to some extent occurred throughout history in sympatric areas. What local factors or ecological relationships that correlate with or trigger hybridization between these species has however been unclear. We studied spatial distribution of hybrids between brown hares and mountain hares in Sweden in relation to characteristics of the sampled localities (hunting grounds). In a sample of 70 brown hares collected from 39 populations in south‐central Sweden during 2003–2005, 11 (16%) showed introgressed mtDNA from mountain hares. Among the brown hares from their northern range, i.e. in general the most recent establishments, the corresponding figure was 75% (9/12). The frequency of samples with hybrid ancestry increased significantly with latitude, altitude and hilliness, and were higher (p<0.1) in recently established populations and/or where the proportion of arable land was low. Several site‐specific parameters were correlated, e.g. latitude as expected to hilliness, and no parameter explained the occurrence of hybrids exclusively. Instead, the appearance of mountain hare mtDNA among brown hares was associated with a conglomerate of parameters reflecting landscapes atypical for the brown hare, e.g. forest dominated and steep areas where the species quite recently was established. We suggest that these abiotic factors mirror the main aspect influencing hybridization frequency, namely the density or relative frequency of the two species. In atypical brown hare landscapes with recent establishment, mountain hares are probably relatively more common. When one species dominate in numbers, or when both species display low densities, increased frequency of hybridization is expected due to low availability of conspecific partners, a phenomenon referred to as Hubbs’ principle.     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

Synergistic influences of introduced herbivores and fire on vegetation change in northern Patagonia,Argentina

Question: We investigated how cattle and European hares, the two most widespread exotic herbivores in Patagonia, affect species composition, life‐form composition and community structure during the first 6 years of vegetation recovery following severe burning of fire‐resistant subalpine forests and fire‐prone tall shrublands. We asked how the effects of introduced herbivores on post‐fire plant community attributes affect flammability of the vegetation. Location: Nahuel Huapi National Park, northwest Patagonia, Argentina Methods: We installed fenced plots to exclude livestock and European hares from severely burned subalpine forests of Nothofagus pumilio and adjacent tall shrublands of N. antarctica. The former is an obligate seed reproducer, whereas the latter and all other woody dominants of the shrubland vigorously resprout after burning. Results: Repeated measures ANOVA of annual measurements over the 2001‐2006 period indicate that cattle and hare exclusion had significant but complex effects on the cover of graminoids, forbs, climber species and woody species in the two burned community types. Significant interactions between the effects of cattle and hares varied by plant life forms between the two communities, which implies that their synergistic effects are community dependent. Conclusions: Following severe fires, the combined effects of cattle and hares inhibit forest recovery and favour transition to shrublands dominated by resprouting woody species. This herbivore‐induced trend in vegetation structure is consistent with the hypothesis that the effects of exotic herbivores at recently burned sites contribute to an increase in the overall flammability of the Patagonian landscape.