Research Article
The arctic fox (Alopex lagopus): an opportunistic specialist
- Bodil Elmhagen, Magnus Tannerfeldt, Paolo Verucci, Anders Angerbjörn
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- 01 June 2000, pp. 139-149
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Reliable and abundant resources are likely to favour specialization, while unpredictable environmental variation should favour a generalist strategy. The rodent population cycles of northern latitudes can be seen as both predictable and unpredictable, depending on the scale in time and space. The arctic fox Alopex lagopus is an opportunistic carnivore, but paradoxically, it seems to function as a specialist on fluctuating rodent Arvicolinae populations in most inland areas. We have studied the dietary response of arctic foxes in Sweden during 5 years of varying abundance of Norwegian lemming Lemmus lemmus, and how these changes influenced the reproductive success of the foxes. The arctic fox population on mainland Fennoscandia is threatened by extinction and the situation has deteriorated during the 1980s and 1990s because of an absence of lemming peaks. Our results showed that in all years, lemming was the main prey for arctic foxes, with 85% frequency of occurrence in summer faeces (scats). Bird remains (mainly Passeriformes) were present in 34% of the scats, reindeer Rangifer tarandus in 21%, voles and shrews in 4% and hares Lepus timidus in 2% of the scats. The occurrences of lemming, bird and larger mammal (reindeer and hare) remains in the scats varied significantly between years. Temporal variations within summer seasons and dietary differences between sub-areas, indicated that arctic foxes fed opportunistically on the alternative prey types. Den occupancy rates were positively correlated with lemming population densities during the previous winter, indicating a strong numerical response. We conclude that from a functional aspect, the arctic fox in Sweden is a lemming specialist, since lemming is the main prey and their abundance is the best predictor of arctic fox reproductive success. Other prey are used opportunistically in relation to their availability.
Social structure of a polygynous tent-making bat, Cynopterus sphinx (Megachiroptera)
- Jay F. Storz, Hari R. Bhat, Thomas H. Kunz
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- 01 June 2000, pp. 151-165
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The social structure of an Old World tent-making bat Cynopterus sphinx (Megachiroptera), was investigated in western India. A combination of census and mark–recapture data over 2 years (1996–98) was used to infer the form of the mating system, compositional stability of social groups and mode of new social group formation. The breeding population of C. sphinx was subdivided into diurnal roosting colonies, each of which contained one to five discrete roosting groups and often one or more solitary bats in adjacent roosts. Bats most frequently roosted in stem tents constructed in the flower/fruit clusters of the kitul palm Caryota urens. Temporal variation in social structure was assessed using visual census data for a subset of the study population over 3 years (1995–98) spanning six consecutive reproductive periods. The sex and age composition of diurnal roosting groups indicated a polygynous harem-forming mode of social organization, as groups invariably contained a single adult male, 1–37 reproductive females and their dependent young (n = 33 harems). Harem size averaged 6.1 adults in the wet season (n = 19, SD = 3.5) and 13.6 adults in the dry season (n = 14, SD> = 8.5). The same harem social configuration was maintained year-round, despite a high degree of synchrony and seasonality in the timing of reproduction. Juveniles of both sexes dispersed after weaning and sexually immature bats were never present in harems at the time of parturition. Adult females often remained associated as roostmates from one parturition period to the next, and group cohesion was unaffected by turnover of harem males. Adult females frequently transferred among roosts within the same colony, and harems underwent periodic fissions and fusions. The founding of new harems most often resulted from the fissioning of previously cohesive harems within the same colony. However, some harems contained disproportionate numbers of yearling females, indicating that new groups are also founded by nulliparous females of the same age cohort. A significant degree of heterogeneity in age composition among harems was revealed in the 1998 dry season, but was unrelated to age-stratification of tent roosts. Although formation of new harems may be non-random with respect to age composition of the founders, founding events are not restricted to newly created tents and often involve recolonization of previously occupied roosts.
Effects of temporal and spatial variations in food supply on the space and habitat use of red squirrels (Sciurus vulgaris L.)
- Peter W. W. Lurz, P. J. Garson, Luc A. Wauters
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- 01 June 2000, pp. 167-178
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In non-native conifer plantations characterized by strong spatial and temporal variations in the availability of tree seeds in Spadeadam Forest, northern England, the home range and habitat use of red squirrels Sciurus vulgaris was very flexible. Males tended to have much larger home ranges than females and core-areas of most breeding females seemed mutually exclusive. Adult female red squirrels were found to increase their home range and core-area size in forest patches where food was less abundant. Home-range size was significantly related to home-range quality and the extent of overlap by other females. In contrast with high-quality continuous conifer forests: (1) a considerable proportion of adult males and females at Spadeadam shifted home range, (2) both sexes had much larger home ranges than reported from other habitats in Britain or Belgium. Many ranges were multinuclear, particularly from January onwards, when supplies of seeds become depleted through consumption and seed shed. Squirrels tracked the availability of conifer seeds (lodgepole pine cones throughout the study, Norway spruce cones in spring 1992 and Sitka spruce cones in autumn 1993) and intensively used several non-adjacent activity centres in temporally food-rich patches. Consequently, habitat preference changed markedly with time. The squirrels seemed to maximize nitrogen intake and to avoid the smaller seeds when possible. This resulted in an overall preference for a mixed diet of lodgepole pine and spruce seeds and avoidance of Sitka spruce seeds when Norway spruce seeds were available. These results lend support to the hypothesis of Ostfeld (1985) that when food is sparse and patchily distributed, females should develop intrasexual territoriality, concentrating their activity in food-rich patches, while males should be non-territorial and adapt their space use to the distribution of females.
Are calcitic layers an effective adaptation against shell dissolution in the Bivalvia?
- E. M. Harper
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- 01 June 2000, pp. 179-186
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Conventional wisdom, based on properties of reagent-grade salts, states that organisms which construct their exoskeletons from calcite rather than aragonite have a greater ability to resist dissolution. However, experiments on individual bivalve microstructures show that other factors such as crystal size and the proportion of organic matrix outweigh constituent mineralogy in determining the rate of shell loss in cold waters. Moreover, the loss of shell material was minimal, corresponding to an annual loss in shell thickness of 3–40 μm depending on microstructure. These findings must cast doubt on the commonly held supposition that calcitic layers added to the outside of shells are an adaptation against dissolution in Holocene ‘Aragonite’ seawater.
Abundance, demographics and body condition of a translocated population of St Lucia whiptail lizards (Cnemidophorus vanzoi)
- H. C. Dickinson, J. E. Fa
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- 01 June 2000, pp. 187-197
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The whiptail lizard Cnemidophorus vanzoi is a large-bodied teiid found only on two islands (Maria Major and Maria Minor), off St Lucia, West Indies. In May 1995, 42 lizards from Maria Major were introduced to the smaller uninhabited Praslin Island on the same coastline. Three years post-release, we studied abundance, demography and morphometrics of the translocated lizard population, during a 6-month period covering wet and dry seasons. Age, sex, snout–vent length (SVL), body mass (BM), tail length, tail regeneration, and overall condition (moulting, reproductive condition, cuts, external parasites) of 107 animals caught during the study are analysed in the present paper. Comparisons are also made with the source population. A body condition index (CI (BM/SVL)), sex ratio (adult males : adult females), age ratio (adults : juveniles), and sexual size dimorphism ratio (SVL adult male : SVL adult females) were calculated for the study population. Distance sampling and mark–re-sight surveys were used to estimate population size and lizard density. A total of 155 ± 26 individuals were estimated. The lizard population was found to have a high growth rate (r = 0.97–3.95). There were significant seasonal changes in lizard abundance. Seasonal differences in lizard numbers, BM and CI suggest either severe resource limitation during the dry season, or selective aestivation. A high frequency of tail autotomy may point to intense intraspecific competition as the island is relatively free from main predators such as the black rat Rattus rattus. Sex ratio, sexual size dimorphism and sexual dichromatism all indicate a territorial species in a generally non-territorial family (Teiidae). Some adult males seem to maintain juvenile colours. It is suggested that the introduced population has successfully colonized its new environment and that no significant change in the animals condition or size has occurred during the 3 years since translocation.
Range extension and microhabitat of Lightiella incisa (Cephalocarida)
- Marleen De Troch, Frank Fiers, Magda Vincx
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- 01 June 2000, pp. 199-204
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During an intensive meiofauna sampling campaign in intertidal seagrass beds along the Caribbean coast of the Yucatán Peninsula (Mexico), 131 specimens of Lightiella incisa (Cephalocarida, Crustacea) were recovered from the sediment. Two-thirds of the specimens were adults, one-third were pre-adults. This collection is the first record of this minute primitive crustacean in the western part of the Caribbean Sea, and extends the known range 3000 km from the type locality of Hastings Bay, Barbados. A detailed sampling protocol and environmental data made it possible to study the microhabitat preferences of this species, and perhaps for cephalocarids in general for the first time. The vertical distribution of L. incisa in the sediment showed a maximum density in deeper layers, i.e. 3–4 and 4–5 cm depth. Nitrate and nitrite concentrations seem to be most closely related to the distribution of L. incisa. It was clear that L. incisa followed polychaetes to deeper sediment layers. In this study we state that L. incisa is an endobenthic species occupying anoxic sediments oxygenated by bioturbation (e.g. Polychaeta) rather than being an animal living in the oxygenated top layers.
Climate, vegetation, and predictable gradients in mammal species richness in southern Africa
- Peter Andrews, Eileen M. O'Brien
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- 01 June 2000, pp. 205-231
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Many hypotheses have been proposed to account for geographic variations in species diversity. In general these relate to some aspect of climate, particularly climatic variables measuring available or potential energy, but while these relate directly to plant diversity they may only indirectly affect mammal species richness. We have examined these relationships by mapping and correlating mammal species richness in southern Africa (n= 285 species) with 15 climatic variables, two topographic variables, and woody plant species richness (n = 1359 species). The effect of area on richness was held a constant by using an equal-area grid cell matrix superimposed on species range maps, with each grid cell equal to 25 000 km2. We found that variability in the plant species richness alone accounts for 75% of the variability in mammal species richness. Of the climatic variables, only thermal seasonality approaches this figure, accounting for 69% of the variability, while annual measures of temperature, precipitation or energy account for only 14–35% of variability. Differences from North American mammal diversity studies, where annual temperature, and hence annual potential evapotranspiration (PET), have been found to be more important, are attributed in part to southern Africa's climate and vegetation being largely temperate to tropical, as opposed to temperate to polar in North America. By distinguishing different types of mammal based on size, spatial and dietary guilds, other differences emerge. Strong correlations with annual temperature exist only for large mammals, accounting for 60–67% of the variability in species richness of large mammals compared with <20% for small mammals. Small mammals are strongly correlated with other climatic or vegetation parameters, especially plant richness and thermal seasonality; frugivorous and insectivorous mammal richness is correlated with thermal seasonality and minimum monthly PET; and arboreal and aerial species richness is correlated with plant richness, thermal seasonality and minimum monthly PET. Up to 77% of the variability in richness of arboreal, frugivorous and insectivorous species can be explained by woody plant richness, compared with only 38–48% of the variability in terrestrial herbivores. It is clear from this that different kinds of mammals are differentially affected by climatic and environmental factors, and this explains some of the discrepancies found in earlier studies where no distinction was made between different sizes or guilds of mammal. This result has implications both for the conservation of mammalian communities at the present time and for understanding the evolution and structure of mammalian communities in the past.
Carnivory in the greater noctule bat (Nyctalus lasiopterus) in Italy
- G. Dondini, S. Vergari
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- 01 June 2000, pp. 233-236
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An analysis of 59 droppings of the greater noctule Nyctalus lasiopterus, collected either during direct handling of the bat or from the bat-boxes used as shelters in the Pian di Novello Natural Reserve (Tuscany, Italy), has shown that it is a partial carnivore. The content of the samples, collected in October and November of 1995 and in September and October of 1996 and 1997, consisted mainly of bird feathers identified as those of the robin Erithacus rubecula and the blue tit Parus caeruleus. The presence of feathers during the 3 consecutive years of the study period confirms that the diet of the greater noctule in this area is not based exclusively on arthropods as was previously suggested.
Fox (Vulpes vulpes) management in three contrasting regions of Britain, in relation to agricultural and sporting interests
- Matthew J. Heydon, Jonathan C. Reynolds
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- 01 June 2000, pp. 237-252
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Fox culling practices in three contrasting rural regions (1280–2320 km2) of Britain were investigated by questionnaire surveys of landowners and tenant farmers. Between 50 and 52% of all farmers identified in each region contributed to the survey (total number of replies = 1123). Fox culling was widespread in all regions, with 70–95% of farmers involved either directly or indirectly as hosts. Motivation for culling foxes and the methods employed reflected regional variation in agricultural and game-shooting interests. Social factors associated with fox-hunting for sport also played an important role in one region. The contribution to the total fox cull made by communally organized groups operating over large tracts of farmland, and the efforts of individual farmers at a local scale, also varied regionally. The numbers of foxes culled were close to published estimates of annual productivity in British fox populations. Deliberate culling was therefore likely to be the chief cause of fox mortality in these regions. The marked differences in culling practices between these regions provide a strong case that the impact of culling on fox population dynamics must be assessed for regions of this size, as well as at a more local scale. Studies at a national scale (e.g. the whole of Britain) are probably of little biological or sociological significance. The success of different interest groups in achieving their aims, and the consequences of any change in culling practices must also be considered at a similar geographical scale.
Variation in abundance of foxes (Vulpes vulpes) between three regions of rural Britain, in relation to landscape and other variables
- Matthew J. Heydon, Jonathan C. Reynolds, Michael J. Short
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- 01 June 2000, pp. 253-264
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Spotlight transect surveys with distance sampling were used to estimate spring (pre-breeding) and autumn (post-production) fox Vulpes vulpes densities in three contrasting rural areas of Britain during 1995–97. This was the first attempt in Britain to measure and compare fox densities over large geographical areas (630–1460 km2). Mean post-production fox abundance was estimated to be 0.90/km2, 2.62/km2, and 0.59/km2 in mid-Wales, the east Midlands and East Anglia, falling to pre-breeding levels of 0.41/km2, 1.17/km2, and 0.16//km2 in spring. As relative measures of regional density, these estimates are strongly supported by independent indices of fox abundance, and by the simultaneous survey of two sympatric species, the badger Meles meles and brown hare Lepus europaeus, which demonstrate the absence of any terrain-related bias. Absolute abundance is less easy to verify, but estimates of spring density based on breeding earth censuses support the transect surveys. For two of the three regions, fox density was close to levels predicted by extrapolation on the basis of landscape, but in the third region (East Anglia), fox density was substantially below prediction. Thus, results failed to support a hypothesis that fox abundance can be predicted solely on the basis of landscape and its close correlates. Rather, they favoured a competing hypothesis that an independent factor determines fox abundance in some regions. A likely factor is culling by man.
Demography of rural foxes (Vulpes vulpes) in relation to cull intensity in three contrasting regions of Britain
- Matthew J. Heydon, Jonathan C. Reynolds
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- 01 June 2000, pp. 265-276
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The impact of deliberate culling of fox populations has been much debated. Although a local (< 10 km2) impact has been accepted, previous authors have denied that culling has any impact on a larger scale because local losses are compensated through immigration. Rather, it has been claimed that at this scale fox density is determined by resources, mediated through social behaviour and breeding suppression. We determined the impact of culling on a regional scale (> 1000 km2), using data on culling (Heydon & Reynolds, 2000), fox density (Heydon, Reynolds & Short, 2000) and productivity. The three U.K. study regions (size 1238–2322 km2) were in mid-Wales (A), the east Midlands (B) and East Anglia (C). High productivity in regions A and C was associated with low density, high culling mortality and high overall mortality (all relative to region B), indicating that density was suppressed by culling. In region B (moderate) breeding suppression was associated with a higher density and lower cull than in regions A and C, implying that fox density was closer to the maximum sustainable by resources. We conclude that the impact of culling in different regions of Britain is variable, dependent on the regional prevalence, methods, and history of culling. However, it is clear that in a range of circumstances culling can substantially depress fox numbers, and that current fox densities reflect a history of culling. This conclusion is fundamental in considering the management of fox predation in farming and conservation contexts.