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.     

Population dynamics of prey exhibiting inducible defenses: the role of associated costs and density-dependence

The effect of antipredator behavior on the dynamics of a resource-consumer model was analyzed in relation to the magnitude of associated costs, and the strength of density-dependence. For this purpose, I present a deterministic continuous resource-consumer model that exhibits biomass conversion, structural homogeneity, and competition for renewable and fixed resources as separate processes. Antipredator behavior is incorporated as an inducible response to consumer density, and has metabolic and feeding costs. By means of numerical methods, I show: (1) that antipredator behavior is stabilizing for certain parameter ranges, where other stabilizing forces do not dominate the dynamics; (2) intraspecific competition for both fixed and renewable resources have a stabilizing role; (3) metabolic cost is always stabilizing, and feeding cost can be stabilizing or destabilizing, depending on the relative strength of the two competition forces.     

Large-scale climatic fluctuation and population dynamics of moose and white-tailed deer

1. Mech et al . (1987) documented cumulative, negative effects of previous winters' snow on rates of population increase in moose ( Alces alces ) and white-tailed deer ( Odocoileus virginianus ), but noted no effect of predation by wolves ( Canis lupus ). Those results were contested by Messier (1991), who analysed smoothed versions of the original abundance data and reported no effect of snow accumulation on population dynamics of either species, but strong effects of wolf predation and food competition.
2. McRoberts, Mech & Peterson (1995) contended that the conclusions reached by Messier (1991) were an artefact of the use of smoothed data. In a subsequent re-analysis of the smoothed data, Messier (1995) argued that the lack of an effect of snow after one year precluded the potential for a cumulative effect beyond one year.
3. We re-analysed original and smoothed data on dynamics of moose and white-tailed deer densities using the same methods as Mech et al . (1987) and Messier (1991), but we used a measure of global climatic fluctuation, the North Atlantic Oscillation (NAO) index. The NAO is the atmospheric process determining most interannual variation in snowfall and winter temperatures in northern latitudes, and its phases drive decadal trends in wintertime precipitation.
4. We observed that rates of increase of moose and white-tailed deer in both the original and smoothed data were influenced by global climatic fluctuation at 2- and 3-year lags, as well as by delayed density-dependent feedback and wolf predation.     

Global climate change and phenotypic variation among red deer cohorts.

The variability of two fitness-related phenotypic traits (body weight and a mandibular skeletal ratio) was analysed among cohorts and age-classes of red deer in Norway. Phenotypic variation among cohorts was pronounced for calves, yearlings and reproductively mature adults. Fluctuations in cohort-specific mean body weights and skeletal ratios of adults correlated with global climatic variation in winter conditions influenced by the North Atlantic Oscillation while cohorts were in utero. Red deer born following warm winters were smaller than those born after cold winters, and this inter-cohort variability persisted into adulthood. Phenotypic variation among cohorts of red deer influenced by climate change may pose consequences for fitness of cohorts since body size and condition contribute to reproductive success and survival in male and female red deer. In particular, the recent trend of increasingly warm winters in northern Europe and Scandinavia may lead to reduced body size and fecundity of red deer, and perhaps other ungulates, in those areas.     

Low genetic variation support bottlenecks in Scandinavian red deer

Loss of genetic variation from genetic drift during population bottlenecks has been shown for many species. Red deer (Cervus elaphus) may have been exposed to bottlenecks due to founder events during postglacial colonisation in the early Holocene and during known population reductions in the eighteenth and nineteenth centuries. In this study, we assess loss of genetic variation in Scandinavian red deer due to potential bottlenecks by comparing microsatellite (n = 14) and mitochondrial DNA variation in the Norwegian and Swedish populations with the Scottish, Lithuanian and Hungarian populations. Bottlenecks are also assessed from the M ratio of populations, heterozygosity excess and from hierarchical Bayesian analyses of their demographic history. Strong genetic drift and differentiation was identified in both Scandinavian populations. Microsatellite variation was lower in both Scandinavian populations compared with the other European populations and mitochondrial DNA variation was especially low in the Swedish population where only one unique haplotype was observed. Loss of microsatellite alleles was demonstrated by low M ratios in all populations except the Hungarian. M ratios’ were especially low in the Scandinavian populations, indicating additional or more severe bottlenecks. Heterozygosity excess compared with the expectation from the number of observed microsatellite alleles suggested a recent bottleneck of low severity in the Norwegian population. Hierarchical Bayesian coalescent analyses consistently yielded estimates of a large ancestral and a small current population size in all investigated European populations and suggested the onset of population decline to be between 5,000 and 10,000 years ago, which coincide well with postglacial colonisation.     

Constraints on plastic responses to climate variation in red deer

Influences of climate on life history traits in natural populations are well documented. However, the implications of between-individual variation in phenotypic plasticity underlying observed trait-environment relationships are rarely considered due to the large, long-term datasets required for such analysis. Studies typically present correlations of annual trait means with climate or assume that individual phenotypic responses are constant. Here, we examine this additional level of variation and show that, in a red deer population on the Isle of Rum, Scotland, changes in climate generate changes in phenotype only amongst individuals who have experienced favourable ecological conditions. Examination of relationships between offspring birth weight and spring temperature within the lifetimes of individual females revealed that the tendency to respond to climate declined as the population density experienced early in life increased. The presence of such systematic variation in individual plasticity is rarely documented in the wild, and has important implications for our understanding of the environmental dependencies of traits under varying ecological conditions.     

Population dynamics of two small pelagic fish in the central-south area off Chile: delayed density-dependence and biological interaction

It is widely believed that environmental variability is the main cause for fluctuations in commercially exploited small pelagic fish populations around the world. Nevertheless, density-dependent factors also can drive population dynamics. In this paper, we analyzed thirteen years of a relative abundance index of two clupeoids fish populations coexisting in the central-south area off Chile, namely the common sardine, Strangomera bentincki, and anchovy, Engraulis ringens. We applied the classical diagnostic tools of time series analysis to the observed time-series. Also, the realized per capita population growth rate was studied with the aim of detecting the feedback structure that is characterizing the population dynamics of the two species. The analysis suggests that population fluctuations of the two species have an important density-dependent component, displaying first-order (direct density-dependent) and second-order (delayed density-dependent) simultaneously. The density-dependent component explained 70.5 and 55.6 % of the realized per capita population growth rate of common sardine and anchovy, respectively. The deterministic skeleton model showed an asymptotic convergence to equilibrium density. In presence of a stochastic environment, fluctuations were reproduced for the species showing a component of fluctuation with a period of 4 year. The intrinsic dynamics of each species is typical of interacting species resulting from trophic interactions. It is postulated that the second-order dynamics of S. bentincki and E. ringens in central-south Chile, may be the result from interactions with a specialist predator (the fishing fleet), interacting with exogenous environmental factors.     

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.     

Population structure, habitat features and genetic structure of managed red deer populations

Management of game ungulates alters population structure and habitat features, with potential effects on genetic structure. Here, we study 26 red deer (Cervus elaphus) populations in Spain. We used census data and habitat features as well as genetic information at 11 microsatellite markers from 717 individuals. We found that metapopulations presented a distribution associated with forest interruptions. Within metapopulations, fences did not have a significant effect on red deer genetic structure. The metapopulations we studied presented similar population structure, but they differed in habitat features and genetic structure. The metapopulation with higher resource availability showed a genetic structure pattern in which genetic relatedness between geographically close individuals was high while relatedness between geographically distant individuals was low. Contrarily, the metapopulation with lower resource availability presented a genetic structure pattern in which the genetic relatedness between individuals of different populations was independent of the geographic distance. We discuss the possible connection between resource availability and genetic structure. Finally, we did not find any population or environmental variable related to genetic differentiation within metapopulations.     

Estimating red deer Cervus elaphus populations: an analysis of variation and cost-effectiveness of counting methods

• 1 Different counting methods are currently used to estimate red deer populations in the open range in Scotland, but there are few data available to compare variation in estimates, or relative cost‐effectiveness.
• 2 While it is impossible to determine the accuracy of counts (as real numbers are unknown), variation within and between different methods can be measured by repeat counts of the same area within as short a period as possible.
• 3 This study aimed to quantify the variation observed from repeat counts using each of four methods (ground, helicopter, infrared helicopter and dung‐counting methods) at one of three study sites in late winters 2003, 2004 and 2005. Additional data from digital camera images of groups from counts in other areas of Scotland were also used to assess the accuracy of visual counts.
• 4 Coefficients of variation (CVs) within any method of between 5% and 16% were recorded, consistent with previous comparisons for red deer open range counts in Scotland. CVs were lowest for ground and helicopter counts. The infrequency of optimal conditions was likely to limit the applicability of infrared counts in Scotland.
• 5 In terms of cost‐effectiveness, helicopter counting was the least labour‐intensive, with costs of other techniques depending on the availability of existing manpower as an overhead cost.
• 6 It is concluded that helicopter counts are most likely to minimize errors while maximizing cost‐efficiency. Accuracy can be improved by the use of digital photography for counting larger deer groups. Estimates are likely to be improved further by increasing the frequency of counts and using the same methods, counters and routes for repeat counts.

     

Liver concentrations of copper, cobalt, and selenium in wild Norwegian red deer (Cervus elaphus)

Liver samples from 245 wild red deer (Cervus elaphus) collected during the licensed hunting season in 2001 from five different locations in western Norway were analyzed for copper (Cu), cobalt (Co), and selenium (Se). The associations between these trace elements and geographical location, age group, and sex were studied. The median (and range of) liver concentrations (microg/g wet weight) for all the examined deer were: Cu 20 (1.7-103), Co 0.08 (<0.01-0.18), and Se 0.09 (0.04-1.0). The results indicate a generally low status of Cu and Se. In total, 15 (6%) red deer had deficient Cu levels (< 4 (microg/g). For all three elements, the liver concentrations showed a significant geographic variation. The geographic difference was most distinct for Cu. The lowest median Cu concentration was found in deer from the island Hitra, where 13% of the animals had deficient Cu levels. Significant differences between age groups were found for all elements, and generally, the adults (> or =2.5 yr) had the highest levels. No significant sex differences within the various age groups were found, with three exceptions: female calves and adults had significantly higher Co levels than male deer, and adult males had significantly higher Se levels than adult females. The Cu and Se status of wild red deer in parts of Norway is low; however, the significance of this needs to be explored further.     

Population dynamics and epidemiology of Toxocara canis in Danish red foxes

Toxocara canis, an intestinal helminth of canids with zoonotic potential, was found in 618 (59%) of 1,040 red foxes (Vulpes vulpes) collected from all Danish provinces (1997-2002). The prevalence and average worm burden were significantly higher for cubs than older foxes and in males than in females. A multiple logistic regression demonstrated that the prevalence was influenced significantly by sex and age of foxes in addition to location, season, and year of collection. The highest prevalence and worm burden were found in rural areas. The size and number of female worms was positively correlated to the fecal egg excretion. The length and fecundity of the worms was significantly higher in male foxes, and a general intensity dependence was suggested from a negative correlation between worm numbers and worm lengths. As compared to intestinal recovery of worms, somatic larvae were recovered from 20% of muscle samples and fecal eggs in 41% of fecal samples. The consistent finding of T. canis larvae in somatic tissues of naturally infected foxes is new.     

Population dynamics and conservation biology of the over-exploited Mediterranean red coral

The main goal of ecologists is nowadays to foster habitat and species conservation. Life-history tables and Leslie-Lewis transition matrices of population growth can be powerful tools suitable for the study of age-structured over harvested and/or endangered species dynamics. Red coral (Corallium rubrum L 1758) is a modular anthozoan endemic to the Mediterranean Sea. This slow growing, long lived species has been harvested since ancient times. In the last decades harvesting pressure increased and the overall Mediterranean yield reduced by 23. Moreover, mass mortality (putatively-linked to global warming) recently affected some coastal populations of this species. Red coral populations are discrete genetic units, gonochoric, composed by several overlapping generations and provided of a discrete (annual) reproduction. A population of this precious octocoral was studied in detail and its static life table was compiled. In order to simulate the trends overtime of the population under different environmental conditions and fishing pressures, a discrete, non-linear model, based on Leslie-Lewis transition matrix, was applied to the demographic data. In this model a bell-shaped curve, based on experimental data, representing the dependence of recruitment on adult colonies density was included. On these bases the stability of the population under different density, reproduction and mortality figures was analysed and simulations of the population trends overtime were set out. Some simulations were also carried out applying to the studied population the mortality values measured during the anomalous mass mortality event which really affected some red coral populations in 1999. The population under study showed high stability and a strong resilience capability, surviving to a 61% reduction of density, to a 27.7% reduction of reproduction rate and to an unselective harvesting affecting 95% of the reproductive colonies.     

Population dynamics of Salmincola salmoneus on Atlantic salmon in a northern Norwegian river

Atlantic salmon Salmo salar are often heavily infected by the gill maggot Salmincola salmoneus, but little information exists on the population dynamics of this parasite. Through a combination of in vivo field examination and laboratory analysis of gills from the Alta River S. salar population in northern Norway, we describe the population dynamics of the parasite and suggest a model for the host-parasite interactions. S. salar did not become infected with S. salmoneus until they returned to the river as first-time spawners. The infection increased rapidly until autumn, and just after spawning 96% of the spent fish (kelts) were infected with a mean intensity of 53 parasites per fish. In May, the prevalence of S. salmoneus on the descending kelts had increased to 100%, but the intensity exhibited little change. A small proportion of the adult S. salar population returned as immature to the river during autumn and had lower parasite intensities than the kelts the following spring. When the fish that had spawned previously (repeat spawners) returned from their second (or more) sea migration, they had an average infection rate of 36 S. salmoneus individuals per fish. The kelts seemed to be the main habitat for the parasite during winter and spring, and they stay long enough in the river to pass the infection to maiden S. salar that enter the river early in summer. These fish then became a source of infection for the maiden fish entering the river later. However, in years that have a possible mismatch between the opposite migration of kelts and maiden S. salar, the immature fall-running and returning repeat spawners will be crucial for maintaining the parasite population. We hypothesize that heavily infected S. salar may suffer reduced growth and survival at sea, potentially reducing the abundance of repeat spawners.     

Population dynamics of Triatoma infestans under natural climatic conditions in the Argentine Chaco

Using experimental chicken houses at a site in central Argentina where the bug Triatoma infestans (Klug) is endemic, nine populations of this vector of Chagas disease were monitored during a 34-month period. Bug populations with four chickens as hosts were consistently larger than those with two chickens as hosts. Age structure of the bug population followed a similar pattern irrespective of the initial age structure. Egg to adult mortality was consistently around 98.5% and there was no consistent evidence for density-dependent mortality. There was some evidence for density dependence in fecundity and recruitment rates, but these were heavily constrained by low temperatures during the winter months. Nymphal development rates correlated most strongly with mean minimum temperatures rather than with mean maximum temperatures. We conclude that vector control using insecticides against this species would be most effective at the onset of winter, when recovery of any surviving populations would be inhibited by low temperatures.