The relative role of winter and spring conditions: linking climate and landscape-scale plant phenology to alpine reindeer body mass

The relative importance of winter harshness and early summer foraging conditions are of prime interest when assessing the effect of global warming on Arctic and mountainous ecosystems. We explored how climate and vegetation onset (satellite-derived normalized difference vegetation index data) determined individual performance in three reindeer populations (data on 27814 calves sampled over 11 years). Snow conditions, spring temperatures and topography were the main determinants of the onset of the vegetation. An earlier onset positively affected the body mass of calves born the following autumn, while there was no significant direct negative impact of the previous winter. This study underlines the major impact of winter and spring climatic conditions, determining the spring and summer food availability, and the subsequent growth of calves among alpine herbivores.     

Dental microwear textural analyses to track feeding ecology of reindeer: a comparison of two contrasting populations in Norway

Using dental microwear textural analysis, we aim to explore variations in feeding habits within and between wild reindeer populations (Rangifer tarandus) in Norway. Specimens from the two wild reindeer areas of Knutshø and Hardangervidda were hunted between late August and the end of September 2014, and found to share very similar topological and ecological characteristics (continental alpine tundra). Despite the homogenisation of plant resources at the end of the snow-free period, dental microwear textural analysis on cheek teeth indicates differences in feeding behaviors between the two populations, in that reindeer from Knutshø might display a more “grazing” signal than the ones from Hardangervidda. Such differences in dental microwear textures reflect the differences in feeding habits that could be linked with differences in population density of reindeer. The Knutshø reindeer density is lower than at Hardangervidda, where primary food resources like grasses, sedges, and lichens are less abundant. Analyses also show the differences in gender for the population of Knutshø, but more studies are necessary for the clearest interpretation of data. Our results emphasize the need to explore local variations at a monthly time frame in order to better assess the versatility of dental microwear textures before applying this kind of analysis to either extant or extinct multi-population scales.     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Functional response in habitat selection and the tradeoffs between foraging niche components in a large herbivore

How herbivore behaviour is influenced by changes in resource levels is central for understanding trophic interactions. We examined whether foraging tradeoffs change with food levels by comparing habitat selection and space use within and between two neighbouring, predator‐free Svalbard reindeer populations. The populations faced different food levels due to contrasting grazing history. Summer resource selection in radiocollared females was assessed by a multi‐dimensional niche approach based on habitat variables obtained from a satellite image (e.g. the normalised difference vegetation index, NDVI) and a digital terrain model. The population at the overgrazed Brøggerhalvøya faced overall lower plant cover, biomass and primary productivity (i.e. lower NDVI) than the population at Sarsøyra. At Brøggerhalvøya, most reindeer selected for productive habitat when choosing home range and patches within the home range. In contrast, habitat selection at Sarsøyra was more affected by abiotic conditions such as moisture, which may influence plant quality. Here, reindeer used patches with even less biomass than the average reindeer at the poorer Brøggerhalvøya. Such a difference in habitat preference with different habitat availability (a functional response in habitat selection) probably reflected increased selection for high‐quality forage at the expense of high forage quantity at Sarsøyra. Accordingly, a negative relationship between habitat productivity and home range size was only present across individuals within Brøggerhalvøya, where forage quantity was the important foraging niche component. Individuals having poor (and large) home ranges apparently could not compensate for this by higher patch selectivity compared to individuals with richer home ranges. The results indicate changes in foraging tradeoffs at contrasting resource levels and that strong interactions occur between habitat selection, space use and the foraging niche structure in the absence of predation.     

The effect of domestic sheep on forage plants of wild reindeer; a landscape scale experiment

Domestic herbivores often compete with wildlife for limited resources, and on longer time-scales, grazing may also increase or decrease coverage of important food plants to wildlife affecting the threshold density for when competition can be expected. In Norway, there are growing concerns about the effect of releasing 2.1 million domestic sheep (Ovis aries) for summer grazing into areas hosting wild populations of alpine reindeer (Rangifer tarandus). We quantified the effect of sheep grazing (0, 25 and 80 sheep/km²) on the development in coverage and abundance of plants known to be important in the diet of reindeer during summer (vascular plants) and winter (lichens) within a fully replicated, landscape scale (2.7 km²) experiment. From 2001 to 2005, the sedge, Carex bigelowii, increased while the herb Solidago virgaurea decreased in frequency at high density of sheep relative to controls (both marginally non-significant). There was no marked development in Deschampsia flexuosa, Salix herbacea or Hierarcium alpinum that could be related to sheep grazing intensity. Lichen coverage and height both decreased at high density of sheep from 2002 to 2005. Effects of low grazing intensity were closer to controls than to high grazing intensity. Our study highlights that high sheep grazing intensity induce changes to the plant community that, at the same time, can improve the summer habitat and detriment the winter habitat to reindeer. Many wild reindeer populations are fragmented and may thus be limited by either summer or winter range. The effect of sheep grazing is predicted to vary accordingly. However, currently, we have limited ability to quantify how much this explicitly means in terms of increased or reduced carrying capacity for reindeer.     

Diet selection by European hares (Lepus europaeus) in the alpine zone of the Snowy Mountains, Australia

European hares (Lepus europaeus) are grazers and open grassland specialists that are replaced in mountain areas of their natural range in the northern hemisphere by browsing/intermediate feeding mountain hares (Lepus timidus), but in their introduced range in the southern hemisphere, occupy the alpine zone. We used micro-histological identification of plant fragments and germination of seeds in faecal pellets of L. europaeus from the Snowy Mountains, Australia, to determine diet. We asked whether diet shifted and/or diet breadth expanded in response to seasonally reduced food availability, particularly during winter. If so, did the constraints of food availability in the alpine zone lead to the diet mirroring that of L. timidus in its native alpine habitat. The diet of L. europaeus was dominated by grasses, herbs and shrubs. The main diet items in summer were grasses (70 %) and herbs (28 %). Grasses declined in the diet between summer and autumn when herbs increased to co-dominance, with a further change after establishment of the winter snowpack to a greater preponderance of shrubs (43 % compared with a maximum of 3 % in snow-free months). L. europaeus selected a wider range of plants in winter (59 species compared with 39 in summer) and diet was significantly more variable in winter than in autumn or summer (and in autumn than summer). We concluded that the persistence of L. europaeus in alpine areas of the southern hemisphere is testament to their ability to expand their dietary breadth to occupy mountain climatic zones normally occupied by L. timidus.     

Food resources of the wild Barbary macaque (Macaca sylvanus) in high-altitude fir forest, Ghomaran Rif, Morocco

A small geographically isolated population of the Barbary macaque inhabits a high-altitude fir forest habitat ( Abies pinsapo ) in the Ghomaran region of the Rif mountains of northern Morocco. The climate of this region is Mediterranean, but the altitude (1600–2100 m) causes winters to be cold (as low as -8.0 C) with snow occurring from November to May (snowfall as deep as 1.5 m). The primary winter feeding adaptation is the ability to ingest high quantities of fir foliage; in spring, the macaques took a high diversity of leafy food items from all vegetation layers; in summer, the macaques foraged terrestrially for a high diversity of food items including seeds, small fruits, bulbous geophytes, and animal foods (including tadpoles from small streams); in autumn, the macaques returned to arboreal foraging, primarily feeding on oak acorns ( Quercus ilex ), fir seeds and yew fruit ( Taxus baccata ). The macaques were capable of ingesting 100 of 195 (51%) of all identified plant species in the region, although during the four-month winter, the macaques only averaged 12.5 common food items. A comparison of the study area with the prime habitat of the Barbary macaque-the high-altitude cedar forests of the Moroccan Moyen Atlas-indicates that climate and vegetation physiognomy are highly similar in both regions. Correspondingly, there is a high degree of similarity in macaque diet in both regions in terms of feeding behaviour by season, food diversity and specific feeding techniques. In the Ghomara, the winter feeding adaptation of fir foliage eating parallels that of the Barbary macaque in cedar forest (winter foraging for cedar foliage). This enables the Barbary macaque to exploit the Ghomaran fir forest habitat during the cold, snowy winters much the same as it does cedar forest habitat throughout a major portion of its geographical range.     

Seasonal dynamics of resource partitioning among foliage-gleaning passerines in northern Finland

Summary The five most abundant species were included in a year-round study with respect to six foraging niche dimensions. Approximately full multidimensional utilization functions were used for niche metrics. During summer foraging overlaps were invariably high, but in other seasons periodically lower. Foraging site breadth was lower in winter, when fewer sites are profitable for foraging than in summer. Feeding posture versatility, by contrast, was highest in winter. Seasonal foraging shifts were very prominent, as great in fact as between-species differences. Often seasonal trends were parallel in different species. Niche axes of macrohabitat-type (e.g. tree) were more open for foraging variation and axes of microhabitat-type (tree part) more rigid. Among the resident species seasonal variation in foraging was greatest in Regulus regulus and Parus cristatus, whereas the foraging behaviour was more stable for P. montanus (an abundant species with broad niche), perhaps owing to greater intraspecific competition. In these northern forests foliage-gleaners must be versatile generalists to cope with the unpredictable resources, and thus they overlap broadly in their general resource niches which are determined by their genetically fixed cost and benefit relations to each resource type. Anyhow, presumably during periodical food shortage, the actual resource uses adjusted by resource availability and competition may overlap narrowly.     

Glucuronidation in the reindeer: dietary modification in the UDP-glucuronosyltransferase activity with 4-nitrophenol, 1-naphthol and phenolphthalein as acceptors

The UDP-glucuronosyltransferase activity towards 4-nitrophenol, 1-naphthol and phenolphthalein was measured from the hepatic microsomes of the reindeer (Rangifer tarandus tarandus) after summer, autumn and winter feeding periods. The microsomes were digested with trypsin or digitonin. The UDP-glucuronosyltransferase activity with 4-nitrophenol and 1-naphthol as aglycones was lower in reindeer on winter food than in ones on summer food after trypsin and digitonin digestion. The activity towards phenolphthalein was the same in each feeding period. The different seasonal feeding affects the structure of microsomal membranes and this is reflected as modifications of the UDP-glucuronosyltransferase towards different substrates.     

Foraging behaviour of Lesser Sheathbills Chionis minor exploiting invertebrates on a sub-Antarctic island

Summary During winter (May through October) many Lesser Sheathbills Chionis minor at Marion Island in the sub-Antarctic were obliged to leave their preferred foraging habitat in penguin colonies to forage for invertebrates on the island's coastal plain. The study describes factors affecting feeding success, time budgets and predation risk of the sheathbills which exploited these small, patchily dispersed prey. The birds appeared to select prey 1 mm in diameter, and ignore smaller, common invertebrates.Sheathbills were highly selective of foraging habitat. During 17 censuses made through the winter, 97% of the 1,504 birdsightings were at only eight of the 19 available vegetation types. Multiple regression analysis revealed that prey density was the most important criterion in habitat preference, followed by plant canopy height and distance of the habitat from the sea. These variables accounted for 78% of the variance of habitat use. Focal-animal observations in a sample of habitats showed that feeding success was correlated with prey density and distance from the sea. Tall vegetation impeded the locomotion and foraging of sheathbills. The sheathbills reduced predation risk from skuas Catharacta lonnbergi and travelling time by foraging near the shore. The spatial distribution of prey within vegetation types was apparently unimportant in habitat selection.During winter 83% of the sheathbills in the study foraged communally and 98% roosted communally. Flocks occurred only on good quality habitat and flocking probably facilitated habitat selection. Feeding success increased initially with increasing flock size but decreased in flocks greater than 15 birds, which was attributed to localized prey deletions. The sheathbills spent 88% of the daytime foraging; and feeding, looking around and walking comprised 99% of foraging time. Feeding time increased with increasing flock size, looking around decreased but walking was unaffected. Aggression was rare, was unaffected by flock size and did not significantly affect feeding. A probability model showed that sheathbills could greatly reduce predation risk by flocking but the benefits would not improve much in flocks greater than eight birds.The habitat selection, time budgets and feeding success of adults, subadults and juveniles were very similar.The exploitation of terrestrial invertebrates by sheathbills was interpreted as an expansion of the population's trophic niche to tap an underexploited resource on a species-poor island.     

Why are shrews so small? The costs and benefits of small size in northern temperateSorex species in the context of foraging habits and prey supply

This paper reviews the ecological advantages and disadvantages of very small body size inSorex Linnaeus, 1758 shrews living at high latitudes with cold winters. It examines the feeding and foraging habits of small and large shrews in the context of prey supply, location of winter prey sources, territory requirements, habitat exploitation and inter-specific competition. Data on feeding habits and prey availability show that the major costs of small size are a reduction in food niche breadth and prey biomass resulting from restrictions on the type and size of prey eaten, and large territory requirements. Major benefits of small size are the ability to subsist on small, numerous and accessible arthropods with high encounter rates, enabling coexistence with larger congeners and exploitation of low-productivity habitats less suitable for larger earthworm-eating species. Small size, coupled with low per capita food intake, is shown to be of special adaptive value in cold winters when food supply is restricted mostly to small arthropods, and earthworms are few.     

Assessing the nutritional consequences of switching foraging behavior in wood bison

Diet is one of the most common traits used to organize species of animals into niches. For ruminant herbivores, the breadth and uniqueness of their dietary niche are placed on a spectrum from browsers that consume woody (i.e., browse) and herbaceous (i.e., forbs) plants, to grazers with graminoid‐rich diets. However, seasonal changes in plant availability and quality can lead to switching of their dietary niche, even within species. In this study, we examined whether a population of wood bison (Bison bison athabascae) in northeast Alberta, Canada, seasonally switched their foraging behavior, and if so, whether this was associated with changes in nutrient acquisition. We hypothesized that bison should switch foraging behaviors from grazing in the winter when standing, dead graminoids are the only foliar plants readily available to browsing during spring and summer as nutritious and digestible foliar parts of browse and forbs become available. If bison are switching foraging strategy to maximize protein consumption, then there should be a corresponding shift in the nutritional niche. Alternatively, if bison are eating different plants, but consuming similar amounts of nutrients, then bison are switching their dietary niche to maintain a particular nutrient composition. We found wood bison were grazers in the winter and spring, but switch to a browsing during summer. However, only winter nutrient consumption of consumed plants differed significantly among seasons. Between spring and summer, bison maintained a specific nutritional composition in their diet despite compositional differences in the consumed plants. Our evidence suggests that bison are selecting plants to maintain a target macronutrient composition. We posit that herbivore''s can and will switch their dietary niche to maintain a target nutrient composition.     

黑龙江省胜山林场冬季驼鹿、马鹿和狍的种间关系

1987—1989年的两个冬季在黑龙江省黑河市胜山林场,对驼鹿、马鹿和狍的食性、采食高度和栖息地进行了研究。驼鹿的主要食物是柳(22.4％)、榛(16.0％)、红松(14.3％)和桦(13.8％);马鹿为杨(36.0％)、桦(22.7％)和红松(17.3％);狍为桦(26.5％)、柳(19.0％)、杨(12.7％)。三种动物采食高度的频次为正态分布。驼鹿的采食高度(标准差)为137.4(±39.0)厘米;马鹿为136.2(±37.0)厘米;狍为84.2(±29.2)厘米。驼鹿和马鹿对红松林的利用程度最高,而狍主要利用沼泽植物和红松林。驼鹿与狍在食物维和栖息地维上的生态位重叠最大,驼鹿与马鹿在采食高度维上的重叠最大。三种动物在食物、采食高度和栖息地三维生态位上的重叠大小的顺序为:驼鹿和马鹿>驼鹿和狍>马鹿和狍。     

Winter habitat–space use in a large arctic herbivore facing contrasting forage abundance

We analysed how changes in resource levels influence foraging trade-offs in late winter by wild Svalbard reindeer. Forage plants, and particularly lichens, were less abundant at the overgrazed Brøggerhalvøya compared with the neighbouring Sarsøyra. Strong interactions occurred between habitat selection, home range size, and feeding crater selection. At Brøggerhalvøya, radiocollared females generally selected productive habitat (high summer NDVI; Normalised Difference Vegetation Index). “Immigrants” at Sarsøyra (dispersed from Brøggerhalvøya in early winter) had similar habitat preferences, probably due to past experience. In contrast, “residents” at Sarsøyra were more influenced by abiotic conditions, using habitat with low NDVI, but selecting for high-quality forage (lichens) when cratering. This suggests more quality-based selection at the expense of quantity when forage abundance increases. Habitat–space use relationships also differed between the animal categories, as home range size decreased with availability of preferred habitat. Thus, changes in forage abundance can strongly influence winter habitat–space use interactions in predator-free systems.     

Dietary overlap of grasshoppers on sandhill rangeland in northeastern Colorado

Summary Overlap in the diets of 14 species of grasshoppers on mixed-grass prairie in northeastern Colorado was estimated by microscopic examination of crop contents. Food availability estimates, taken by a weight-estimate method, facilitated the determination of food preferences. Two cases of high overlap in diets and period of habitat occupancy occurred between species which consumed equal or almost equal numbers of different foods (foodniche breadth) and had almost identical foraging behavior. The first case occurred early in the season between Xanthippus corallipes and Arphia conspersa with foodniche breadths of 14 and 11, respectively. The second case occurred later in the season between Amphitornus coloradus and Trachyrhachys kiowa both of which have foodniche breadths of 6. The amount of movement between plants while feeding was equal or almost equal in both cases, thus we observed high dietary overlap between species with equal ecological efficiencies. Dietary overlap between the 14 grasshopper species decreased as foodniche dimensions increased and the grasshopper population as a whole utilized each food resource approximately proportional to its relative abundance. Factors influencing the interpretation and importance of dietary overlap in this study included 1. non-synchronous life cycles, 2. seasonal changes in diets and food preferences, 3. complexity of the food resources, 4. variations in foodniche dimensions among grasshopper species, and 5. variations in diets between males and females within a species. Our findings agree with MacArthur and Levins theory that the number of competing species which can coexist is proportional to the total range of the environment divided by the niche breadth of the species.     

城市公园植被特征对陆生鸟类集团的影响

城市公共绿地是城市生态系统中重要的鸟类栖息地,其植被特征对鸟类集团存在显著影响。在通过分析植被特征对陆生鸟类集团的作用,从而为公园合理配置植被来提高其作为野生动物栖息地的生态服务功能提供理论基础。2009年10月至2011年10月,采用样线法对上海滨江森林公园进行鸟类调查,利用主成分分析划分鸟类集团,用高度定义植被层次,用卡方检验分析鸟类行为在植被层次上的差异。结果表明,滨江森林公园陆生鸟类群落在乔木层的栖息行为和运动行为频次显著多于其在灌木层和地被层的行为频次,在地被层的取食行为频次显著多于其在乔木层和灌木层的行为频次。陆生鸟类可划分为8个鸟类集团,鸟类集团之间存在栖息、运动和取食空间生态位的重叠。食虫拾取集团、杂食拾取集团、食肉飞取集团和植食拾取集团在栖息、运动和取食空间生态位上均存在较高的重叠度,其通过食性分离各自空间生态位。食虫探取集团和食虫飞取集团互为栖息空间生态位重叠度最高集团,其通过取食方式的不同来实现生态位的分离。根据公园植被特征对鸟类集团的影响结果对上海市公园绿地植被配置提出了建议。     

Experimental evidence of density dependent activity pattern of a large herbivore in an alpine ecosystem

Density dependent processes affecting foraging strategies may in turn influence vital rates and population regulation in large herbivores. Increased competition may lower both forage availability and quality, but whether the main activity constraint at high density is increased searching time or increased digestion time is poorly investigated. In a fully replicated landscape‐scale experiment, we used long‐term data (2003–2009) from domestic sheep grazing at high and low density (80 and 25 sheep km^–2, respectively) on alpine summer ranges to test density dependence in allocation of time to feeding (moving) vs digestion (resting) activities and how this in turn affected body growth. Sheep at high density spent more time actively feeding than sheep at low density, but sheep moved shorter distances while foraging at high density. Increased activity levels at high density suggest that the main activity constraint at high density was availability of high‐quality food increasing searching time and possibly reducing intake rates. Increased movement distances at low density is consistent with a higher selection for more productive vegetation types since high‐quality patches are dispersed in the landscape. The alternative hypothesis, that food processing time increased at high density was not supported as it would have reduced overall activity levels. Individual activity levels increased body growth, but this was not sufficient to fully compensate for lower habitat quality leading to an overall reduced body growth at high density. Our experiment clearly documents changes in activity budgets and movement distances of a large herbivore at high population density, providing one potential behavioural mechanism of density dependent responses observed in vital rates.     

The effects of extreme seasonality of climate and day length on the activity budget and diet of semi‐commensal chacma baboons (Papio ursinus) in the Cape Peninsula of South Africa

We examined the effects of extreme seasonality on the activity budget and diet of wild chacma baboons with access to a high‐quality, human‐derived food source. The Cape Peninsula of South Africa is unusual among nonhuman primate habitats due to its seasonal extremes in day length and climate. Winter days are markedly shorter and colder than summer days but have higher rainfall and higher primary production of annually flowering plants. This combination of fewer daylight hours but higher rainfall is substantially different from the ecological constraints faced by both equatorial baboon populations and those living in temperate climates with summer rainfall. We sought to understand how these seasonal differences affect time budgets of food‐enhanced troops in comparison to both other food‐enhanced troops and wild foraging troops at similar latitudes. Our results revealed significant seasonal differences in activity budget and diet, a finding that contrasts with other baboon populations with access to high‐return anthropogenic foods. Similar to nonprovisioned troops at similar latitudes, troop members spent more time feeding, socializing, and traveling during the long summer days compared to the short winter days, and proportionately more time feeding and less time resting in summer compared to winter. Summer diets consisted mainly of fynbos and nonindigenous foods, whereas winter diets were dominated by annually flowering plants (mainly grasses) and ostrich pellets raided from a nearby ostrich farm. In this case, food enhancement may have effectively exaggerated seasonal differences in activity budgets by providing access to a high‐return food (ostrich pellets) that was spatially and temporally coincident with abundant winter fallback foods (grasses). The frequent use of both alien vegetation and high‐return, human‐derived foods highlights the dietary flexibility of baboons as a key element of their overall success in rapidly transforming environments such as the South African Cape Peninsula. Am. J. Primatol. 72:104–112, 2010. © 2009 Wiley‐Liss, Inc.     

Effects of habitat fragmentation on foraging behaviour of tits and related species: does niche space vary in relation to size and degree of isolation of forest fragments?

We studied the winter foraging niches of tits and related species in deciduous forest fragments varying in size between 1 and 30 ha (plus one forest of 200 ha) in order to investigate the influence of forest fragmentation on foraging niches Very few correlations between niche structure (foraging niche, width and overlap) and forest size or isolation turned out to be significant This implies that either the species that disappear in small fragments are those that suffer most from competition (making the effect unmeasurable), or that competition is relatively unimportant for niche structure In any case we find no evidence that foraging niches are strongly affected by the changes (in habitat and/or community structure) associated with fragmentation