Foraging conditions, tooth wear and herbivore body reserves: a study of female reindeer

Several aspects of the life history of reindeer (Rangifer tarandus) are related to the nutritional condition of the animals. Moreover, compensatory growth and fattening in summer decreases with age. The interaction of tooth wear and the standing crop of lichens on age-related variation in body size and tissue reserves was examined to evaluate the proximate causes of density-dependent food limitation on life history parameters in female reindeer. Studies in nine semi-domesticated free-ranging reindeer herds showed that molar height depended on the mean standing crop of terrestrial lichens in winter habitats. The extent of tooth wear had the strongest effect on body reserves among the oldest females (11–14 years). This indicates that severe tooth wear limits the animals′ ability to process food efficiently and, hence, to maintain their body reserves. Both tooth wear and the biomass of lichens influenced body mass in old females, probably because on heavily exploited winter ranges reindeer had to use higher proportions of lower-ranking coarser foods, especially dwarf shrubs. Received: 1 December 1997 / Accepted: 13 July 1998     

Environmental and Physiological Influences to Isotopic Ratios of N and Protein Status in a Montane Ungulate in Winter

Winter severity can influence large herbivore populations through a reduction in maternal proteins available for reproduction. Nitrogen (N) isotopes in blood fractions can be used to track the use of body proteins in northern and montane ungulates. We studied 113 adult female caribou for 13 years throughout a series of severe winters that reduced population size and offspring mass. After these severe winters, offspring mass increased but the size of the population remained low. We devised a conceptual model for routing of isotopic N in blood in the context of the severe environmental conditions experienced by this population. We measured δ¹⁵N in three blood fractions and predicted the relative mobilization of dietary and body proteins. The δ ¹⁵N of the body protein pool varied by 4‰ and 46% of the variance was associated with year. Annual variation in δ¹⁵N of body protein likely reflected the fall/early winter diet and winter locations, yet 15% of the isotopic variation in amino acid N was due to body proteins. Consistent isotopic differences among blood N pools indicated that animals tolerated fluxes in diet and body stores. Conservation of body protein in caribou is the result of active exchange among diet and body N pools. Adult females were robust to historically severe winter conditions and prioritized body condition and survival over early investment in offspring. For a vagile ungulate residing at low densities in a predator-rich environment, protein restrictions in winter may not be the primary limiting factor for reproduction.     

Dynamics of a harvested moose population in a variable environment

1. Population size, calves per female, female mean age and adult sex ratio of a moose ( Alces alces ) population in Vefsn, northern Norway were reconstructed from 1967 to 1993 using cohort analysis and catch-at-age data from 96% (6752) of all individuals harvested.
2. The dynamics of the population were influenced mainly by density-dependent harvesting, stochastic variation in climate and intrinsic variation in the age-structure of the female segment of the population.
3. A time delay in the assignment of hunting permits in relation to population size increased fluctuations in population size.
4. Selective harvesting of calves and yearlings increased the mean age of adult females in the population, and, because fecundity in moose is strongly age-specific, the number of calves per female concordantly increased. However, after years with high recruitment, the adult mean age decreased as large cohorts entered the adult age-groups. This age-structure effect generated cycles in the rate of recruitment to the population and fluctuations introduced time-lags in the population dynamics.
5. An inverse relationship between recruitment rate and population density, mediated by a density-dependent decrease in female body condition, could potentially have constituted a regulatory mechanism in the dynamics of the population, but this effect was counteracted by a density-dependent increase in the mean age of adult females.
6. Stochastic variation in winter snow depth and summer temperature had delayed effects on recruitment rate and in turn population growth rate, apparently through effects on female body condition before conception.     

Survival of Adult Martens in Northern Wisconsin

Abstract: Low adult marten (Martes americana) survival may be one factor limiting their population growth >30 yr after their reintroduction in Wisconsin, USA. We estimated annual adult marten survival at 0.81 in northern Wisconsin, with lower survival during winter (0.87) than summer-fall (1.00). Fisher (Martes pennanti) and raptor kills were infrequent, and each reduced marten adult annual survival <10%. Annual adult survival was similar to or higher than survival in other areas, suggesting that it was not unusually low and therefore did not limit recovery of marten populations in northern Wisconsin. We captured few juvenile martens, suggesting low reproduction or reduced juvenile survival.     

Decelerating and sex-dependent tooth wear in Norwegian red deer

In ungulates, tooth wear is often suggested as a proximate cause of senescence. Tooth wear is expected to be sex-dependent since energetic requirements and food selection varies largely between sexes in sexually dimorphic ungulates. Furthermore, tooth wear may lower mastication efficiency, and we predict a negative correlation between tooth wear and body weight or condition. We tested these predictions on data on tooth wear (estimated as height of first molar) of 1,311 male and 1,348 female red deer ( Cervus elaphus) aged 3-25 years and harvested along the west coast of Norway. The rate of tooth wear decreased with age. Males wear teeth at a higher rate (from 0.61 mm/year in 4-year olds to 0.45 mm/year in 11-year olds) than females (from 0.52 mm/year in 4-year olds to 0.39 mm/year in 11-year olds). Molar height correlated positively with body weight in both sexes, but not after adjusting for body size. Molar height was strongly dependent on body size in 3-year-old individuals (when tooth wear is minimal). Earlier reports in the literature of a positive correlation between tooth height and body weight may therefore be due to initial size differences rather than differences in condition due to tooth wear.     

Short- and long-term effects of winter and spring weather on growth and survival of red deer in Norway

Populations of red deer (Cervus elaphus) in Norway have increased continuously over the last decades. We tested the possible effects of climate and increase in population size on the survival rates and body condition of individuals in one of the northernmost populations of red deer in Europe. Based on 678 individuals of known age marked between 1977 and 1995, we estimated annual survival rates, the probabilities of being harvested and the recapture probability according to sex, age, year, winter and spring weather, population size, and, body weight and body condition, using capture-mark-recapture models. Winter harshness negatively influenced the body weight of yearlings and the survival of calves of both sexes. Spring weather influenced the survival of males in all age classes. A negative trend during the study period was detected in body weight and condition of calves and yearlings, but not in any age- or sex- specific survival rates. No significant gender differences in mean survival were shown in any age class. Moreover, there was little (male) or no (female) detectable between-year variation in survival rates for yearlings and adults. Winter weather acts as a limiting factor on population growth through a short-term effect on first-year survival and a long-term effect on body weight. We discuss the surprising low sex differences in natural survival rates and the differential effects of winter harshness on body weight, body condition and survival in relation to life history characteristics of red deer. Received: 10 November 1997 / Accepted: 2 June 1998     

Variation in body size and metamorphic traits of Iberian spadefoot toads over a short geographic distance

Determinants of geographic variation in body size are often poorly understood, especially in organisms with complex life cycles. We examined patterns of adult body size and metamorphic traits variation in Iberian spadefoot toad ( Pelobates cultripes ) populations, which exhibit an extreme reduction in adult body size, 71.6% reduction in body mass, within just about 30 km at south-western Spain. We hypothesized that size at and time to metamorphosis would be predictive of the spatial pattern observed in adult body size. Larvae from eight populations were raised in a common garden experiment at two different larval densities that allow to differentiate whether population divergence was genetically based or was simply a reflection of environmental variation and, in addition, whether this population divergence was modulated by differing crowding larval environments. Larger adult size populations had higher larval growth rates, attaining larger sizes at metamorphosis, and exhibited higher survival than smaller-sized populations at both densities, although accentuated at a low larval density. These population differences appeared to be consistent once embryo size variation was controlled for, suggesting that this phenotypic divergence is not due to maternal effects. Our results suggest considerable genetic differentiation in metamorphic traits that parallels and may be a causal determinant of geographic variation in adult body size.     

A critique of the «increasing population density effect»

The «increasing population density effect» (IPDE) proposed by Macchiarelli and Bondioli (1986) represents one of the most recent attempts to account for dental reduction in modern human populations. Under the IPDE, the marked reduction in tooth size observed in post-Pleistocene human groups is seen as merely a side-effect of a more general reduction in body size, which resulted from an increase in population density. The model is dependent upon a strong correlation between tooth size and body size, which after numerous attempts has yet to be convincingly demonstrated in humans. This paper argues that the IPDE neglects the negative consequences on individual fitness of teeth which are too large to fit into diminishing jaws or are more susceptible to pathology, and that the worldwide reduction of tooth size is the result of selection for smaller teeth due to shifts to a softer and/or more cariogenic diet. Although increased population density may have intensified this selective effect by decreasing general fitness by lowering resistance to oral infections, it was not the primary cause of dental reduction. All proposed mechanisms of dental reduction, however, are in need of additional testing, and possible directions are offered for future studies of the phenomenon.     

Sea ice affects the population dynamics of Adélie penguins in Terre Adélie

Overall Adélie penguin population size in Pointe Géologie Archipelago increased between 1984 and 2003 at a rate of 1.77% per year, and averaged 33,726±5,867 pairs. As predicted by the optimum model proposed by Smith et al. (Bioscience 49:393–404, 1999). Adélie penguin population size increased when sea ice extent and concentration (SIE and SIC) decreased six years earlier, indicating that the conditions around reproduction or first years at sea, were determinant. The breeding success averaged 85.2±35.45% and was not related to environmental variables. Adult survival probability varied between years from 0.64 to 0.82. Southern oscillation index (SOI) had a strong negative effect on adult annual survival. Adult survival of Adélie penguins increased during warmer events, especially during winter and spring at the beginning of reproduction. Therefore, we speculate that the rapid decreases in 1988–1991 and 1996 of the breeding population size were related to a decrease in adult mortality. However, adult survival varied little, and could not explain the strong increasing population trend. The sea ice conditions during breeding or during the first year at sea appeared determinant and influenced the population dynamics through cohort effects, probably related to the availability of productive feeding habitats.     

Philopatric predisposition to predation-induced ecological traps: habitat-dependent mortality of breeding eiders

Because population size is sensitive to changes in adult survival, adult survival may be buffered against environmental variability. Philopatry may be adaptive in changing environments, but it could also constrain breeding habitat selection under changing conditions such as shifting predation regimes. Habitat preference and quality could become decoupled in long-lived philopatric species that evolved in stable environments when suddenly faced by increased adult predation risk, as dispersal may be triggered by past reproductive failure. We evaluated whether the Baltic eider (Somateria m. mollissima) population may currently face a predation-induced ecological trap. Eiders are philopatric and nest on open and forested islands. We hypothesized that open-nesting females would be disproportionately affected by increased predation. We compared female annual survival in these two habitats in 1996–2010. We also tested for effects of time trends, winter severity (NAO), female body condition, and habitat-specific predation pressure on survival. Our results revealed the lowest survival recorded for this species (Φ?=?0.720), and survival on open islands was significantly lower (Φ?=?0.679) than on forested islands (Φ?=?0.761). Nonetheless, only 0.7?% of females changed breeding habitat type despite ample availability of alternative islands, and breeding phenology in both habitats was similar. Female survival increased with body condition, while it was unrelated to winter climate and stable over time. Open islands had a higher predation pressure on incubating females. Breeding philopatry results in a predator-mediated ecological trap for open-nesting eiders. Our results contribute to explaining the drastic decline of the Baltic eider population.     

Population variables and causes of mortality in a hedgehog (Erinaceous europaeus) population in southern Sweden

A hedgehog population in southern Sweden was studied by capture-mark-recapture during 1972–1979. Each animal was weighed, sexed and marked with one numbered clip in each ear.
The number of subadult and adult hedgehogs was about 20 in the first two years, 1972–1973. Numbers then increased and approximately doubled and the population peaked in 1978. A decrease came in 1979.
The estimated number of juveniles produced varied between years with the lowest figure (15 juv.) in 1974 when the population was low and the highest in 1977 (39 juv.) when the population was high. As no captured subadult female had suckled young, estimates of recruitment rate assumed that females did not reproduce until they were adult. The average number of juveniles per adult female was 2.8. Recruitment rate did not show any correlation with non-juvenile population densities.
For both juveniles and non-juveniles the annual mortality varied greatly between years. Average yearly mortality was 47% for non-juveniles and 34% for juveniles.
For non-juveniles, most of the total mortality occurred during winter, ranging between 26% and 43% (average 33%), and increased with increasing densities of non juveniles. Juvenile winter mortality varied between 6% and 94%, averaging 33%.
Summer mortality of non-juveniles averaged 15%, ranging between 3% and 22%. Traffic kills were the predominant cause of death. Only a few juveniles were found dead during summer and autumn.
There were no statistical differences in age-specific survival rate between males and females and, generally, no differences in survival rate between the age categories within each sex. It is argued that the population size appears to be more influenced by environmental factors, such as food availability, winter nest sites and winter climate, than by density-dependent factors.     

Low larval densities in northern populations reinforce range expansion by a Mediterranean damselfly

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Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

The ecology and evolution of tooth wear in red deer and moose

Differences in body size and diet type (browser–grazer continuum) have formed functional traits of ruminants, including tooth design. Grazers and mixed-feeders eat a more fibrous diet than browsers, which arguably increase tooth wear. Tooth wear has also been suggested to increase with body size. Moreover, for species with large distribution ranges, different populations may be exposed to very different ecological factors affecting diet and thus tooth wear rates. Therefore, evolutionary history and contemporary ecological conditions, both operating through diet, may be important for patterns of tooth wear. Here, we compare inter- and intraspecific rates of tooth wear in multiple populations of one large browser (moose Alces alces ) and one mixed-feeder (red deer Cervus elaphus ) covering the main distribution range of each species in Norway. We found that the mixed-feeding red deer wore teeth faster than the larger and browsing moose, suggesting that feeding-type was more important than body size for patterns of wear. There was substantial spatial variation in tooth wear rates, but the inter-specific difference in wear was consistent. Molar wear rates, but not incisors wear rates, in the browser were less variable between populations than in the mixed-feeder. There was no close link between incisor and molar wear rates at the population level. Our findings are consistent with the view that both evolution related to diet type and current ecological conditions (being proxies for within-species variation in diet quality) are important for patterns of tooth wear.     

Global climate change and reindeer: effects of winter weather on the autumn weight and growth of calves

Reindeer/caribou (Rangifer tarandus), which constitute a biological resource of vital importance for the physical and cultural survival of Arctic residents, and inhabit extremely seasonal environments, have received little attention in the global change debate. We investigated how body weight and growth rate of reindeer calves were affected by large-scale climatic variability [measured by the North Atlantic Oscillation (NAO) winter index] and density in one population in central Norway. Body weights of calves in summer and early winter, as well as their growth rate (summer to early winter), were significantly influenced by density and the NAO index when cohorts were in utero. Males were heavier and had higher absolute growth than females, but there was no evidence that preweaning condition of male and female calves were influenced differently by the NAO winter index. Increasing NAO index had a negative effect on calves' body weight and growth rate. Increasing density significantly reduced body weight and growth rate of calves, and accentuated the effect of the NAO winter index. Winters with a higher NAO index are thus severe for reindeer calves in this area and their effects are associated with nutritional stress experienced by the dams during pregnancy or immediately after calving. Moreover, increased density may enhance intra-specific competition and limits food available at the individual level within cohorts. We conclude that if the current pattern of global warming continues, with greater change occurring in northern latitudes and during winter as is predicted, reduced body weight of reindeer calves may be a consequence in areas where winters with a high NAO index are severe. This will likely have an effect on the livelihood of many northern indigenous peoples, both economically and culturally.     

Adult survival and reproductive rate are linked to habitat preference in territorial,year‐round resident Song Sparrows Melospiza melodia

Individual animal fitness can be strongly influenced by the ability to recognize habitat features which may be beneficial. Many studies focus on the effects of habitat on annual reproductive rate, even though adult survival is typically a greater influence on fitness and population growth in vertebrate species with intermediate to long lifespans. Understanding the effects of preferred habitat on individuals over the annual cycle is therefore necessary to predict its influences on individual fitness. This is particularly true in species that are resident and territorial year‐round in the temperate zone, which may face potential trade‐offs between habitat that maximizes reproduction and that which maximizes non‐breeding season (‘over‐winter’) survival. We used a 37‐year study of Song Sparrows Melospiza melodia residing territorially year‐round on a small island to examine what habitat features influenced adult over‐winter survival, how site‐specific variation in adult survival vs. annual reproductive rate influenced long‐term habitat preference, and if preferred sites on average conferred higher individual fitness. Habitat features such as area of shrub cover and exposure to intertidal coastline predicted adult over‐winter survival independent of individual age or sex, population size, or winter weather. Long‐term habitat preference (measured as occupation rate) was better predicted by site‐specific annual reproductive rate than by expected over‐winter survival, but preferred sites maximized fitness on average over the entire annual cycle,. Although adult over‐winter survival had a greater influence on population growth (λ) than did reproductive rate, the influence of reproductive rate on λ increased in preferred sites because site‐specific variation in reproductive rate was higher than variation in expected over‐winter survival. Because preferred habitats tended to have higher mean site‐specific reproductive and adult survival rates, territorial birds in this population do not appear to experience seasonal trade‐offs in preferred habitat but are predicted to incur substantial fitness costs of settling in less‐preferred sites.     

Survival of British Sedge Warblers Acrocephalus schoenobaenus in relation to west African rainfall

Fluctuations in the population levels and annual adult survival rates of British Sedge Warblers Acrocephalus schoenobaenus since the late 1960s are strongly correlated with indices of wet season rainfall in the west African winter quarters. Population changes are unrelated to estimates of breeding productivity in the previous year. Habitat availability in the winter quarters has probably been the main factor limiting the size of the Sedge Warbler population in Britain during the period of study.     

臭鼩的繁殖和种群年龄结构

对浙江萧山瓜沥区农田臭鼩的研究,发现种群性比不等,存在明显的两性异型。分析了雄体的性成熟率和雌体的怀孕率及性活动率的变化规律。一年中存在两个繁殖高峰期。以齿长指数为指标．臭鼩种群可划分为4个年龄组。探讨了种群年龄结构的季节变化及其自然寿命。     

Catch-up growth in Japanese quail (Coturnix Japonica): relationships with food intake, metabolic rate and sex

The effects of early environmental conditions can profoundly affect individual development and adult phenotype. In birds, limiting resources can affect growth as nestlings, but also fitness and survival as adults. Following periods of food restriction, individuals may accelerate development, undergoing a period of rapid “catch-up” growth, in an attempt to reach the appropriate size at adulthood. Previous studies of altricial birds have shown that catch-up growth can have negative consequences in adulthood, although this has not been explored in species with different developmental strategies. Here, we investigated the effects of resource limitation and the subsequent period of catch-up growth, on the morphological and metabolic phenotype of adult Japanese quail (Coturnix japonica), a species with a precocial developmental strategy. Because males and females differ in adult body size, we also test whether food restriction had sex-specific effects. Birds that underwent food restriction early in development had muscles of similar size and functional maturity, but lower adult body mass than controls. There was no evidence of sex-specific sensitivity of food restriction on adult body mass; however, there was evidence for body size. Females fed ad lib were larger than males fed ad lib, while females subjected to food restriction were of similar size to males. Adults that had previously experienced food restriction did not have an elevated metabolic rate, suggesting that in contrast to altricial nestlings, there was no metabolic carry-over effect of catch-up growth into adulthood. While Japanese quail can undergo accelerated growth after re-feeding, timing of food restriction may be important to adult size, particularly in females. However, greater developmental flexibility compared to altricial birds may contribute to the lack of metabolic carryover effects at adulthood.     

Effects of age, sex and density on body weight of Norwegian red deer: evidence of density-dependent senescence

There are only a few recent studies that have demonstrated senescence in ungulates and nothing is known regarding how patterns of senescence may vary as a function of density Senescence in general is linked to the cost of reproduction, which probably increases with density in ungulates and may differ between the sexes. Further, senescence in ungulates is also regarded to be a function of tooth wear rates. As density dependence and sexual differences in food choice have been well documented, this may lead to different tooth wear rates and, thus, possibly density-dependent and sex-specific patterns of senescence. We therefore investigated the effects of age, sex, density and their possible interactions on the variability of body weight in 29,047 red deer harvested during 1965-1998 from Norway, out of which 380 males and 1452 females were eight years or older. There was clear evidence that spatio-temporal variation in density correlated negatively with body weights. In addition, there was evidence of senescence in both male and female red deer. Age at onset of senescence in females was after 20 years of age and independent of population density. In males, the onset and rate of senescence increased with increasing population density. The onset of senescence for males was at ca. 12 years of age at low density, but decreased to approximately ten years of age at high density. The pattern of density-dependent senescence in males, but not that in females, can be explained if (i) the cost of reproduction increases with density more strongly in male than in female red deer, and/or (ii) tooth wear rates are density dependent in males, but not in females. We discuss the ability of these two different, not mutually exclusive hypotheses in explaining the observed pattern of senescence.