Lineage Loss in Serengeti Cheetahs: Consequences of High Reproductive Variance and Heritability of Fitness on Effective Population Size

Abstract: In natural populations, many breeders do not leave surviving offspring, and as a result many potential genetic lineages are lost. I examined lineage extinction in Serengeti cheetahs ( Acinonyx jubatus ) and found that 76% of matrilines were lost over a 25-year period. Production of future breeders was nonrandom and generally confined to a few families. Five out of 63 matrilines accounted for 45% of the total cheetah population over the course of the study. Lineage persistence is perhaps best illustrated by the variance in lifetime reproductive success ( LRS) and heritability in this parameter. In female cheetahs, variance in LRS was high, and new data show that this LRS was heritable. Variance in LRS and heritability in LRS have dramatic consequences for effective population size, N _e. I calculated N _e for cheetahs, taking into account fluctuating population size, unequal sex ratio, non-Poisson distribution of reproductive success, and heritability of fitness. The N _e was most strongly affected by variance in reproductive success and especially heritability in reproductive success. The variance N _e was 44% of the actual population size, and the inclusion of heritability further reduced N _e to only 15% of the actual population, a ratio similar to that of a social carnivore with reproductive suppression. The current cheetah population in the Serengeti is below numbers suggested by N _e estimates as sufficient to maintain sufficient genetic diversity.     

冶勒自然保护区林麝对生境选择研究

1995年在相岭山系的冕宁冶勒自然保护区，采用作者新近提出的临时空间样方法，以Vanderloeg和Scavia选择指数对林麝对生境的选择进行了研究，结果表明7～8月份，林麝喜欢活动于竹子密度不大（n＝0～15m     

Habitat and population modelling of roe deer using an interactive geographic information system

Management of German roe deer (Capreolus capreolus) populations is a challenge for wildlife managers and foresters because population densities are difficult to estimate in forests and forest regeneration can be negatively affected when roe deer density is high. We describe a model to determine deer population densities compatible with forest management goals, and to assess harvest rates necessary to maintain desired deer densities. A geographic information system (GIS) was used to model wildlife habitat and population dynamics over time. Our model interactively incorporates knowledge of field biologists and foresters via a graphical user interface (GUI). Calibration of the model with deer damage maps allowed us to evaluate density dependence of a roe deer population. Incorporation of local knowledge into temporally dynamic and spatial models increases understanding of population dynamics and improves wildlife management.     

Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Fitness consequences of dispersal decisions for male mountain gorillas (Gorilla beringei beringei)

The fitness consequences of dispersal decisions are difficult to quantify, especially for long-lived species with complex social systems. To calculate those consequences for male mountain gorillas, from the perspective of both subordinate and dominant males, we used behavioral and demographic data obtained over 30 years from the Virunga Volcano population to develop an agent-based model that simulates the life history events, social structure, and population dynamics of the species. The model included variables for birth rates, mortality rates, dispersal patterns, and reproductive skew. The model predicted an average lifetime reproductive success (LRS) of 3.2 for philopatric males (followers) and 1.6 for emigrants. The benefits of philopatry were most sensitive to opportunities for social queuing and to female transfer preferences, but philopatry remained the best strategy over a wide range of group conditions and hypothetical simulations. The average LRS for dominant males was 4.5 when a subordinate stayed and 4.6 when the subordinate emigrated. The dispersal decision of the subordinate male had little impact on the fitness of the dominant male because it came relatively late in the dominant males reproductive life span, and it changed his group composition only incrementally. The fitness consequences for the dominant male were most sensitive to the degree of reproductive skew. Since subordinates suffer a fitness loss when they leave a group, they should accept whatever reproductive restraint is needed to avoid eviction, and the dominant male does not need to offer concessions for them to stay. The dominant male may offer reproductive concessions for other reasons, such as peace incentives or to confuse paternity, or he may not have complete control of reproduction within his group.Communicated by D. Watts     

Polychlorinated biphenyl (PCB) residues in European roe deer (Capreolus capreolus) and red deer (Cervus elaphus) from north-western Poland

The purpose of this study was to evaluate polychlorinated biphenyls (PCBs) contamination levels in roe and red deer from north-western Poland and to assess environmental pollution in this area. A quantitative analysis was conducted using a capillary gas chromatography/mass spectrometry method. The mean concentrations of ΣPCBs (sum of PCBs: 28, 52, 101, 138, 153, 180) in liver samples were 30.24±12.35 ng·g^?1 of lipid weight (l.w.) in roe deer and 60.13±14.23 ng·g^?1 l.w. in red deer, compared with 24.21±10.02 and 45.22±9.77 ng·g^?1 in the lungs of roe and red deer, respectively. PCBs 138, 153 and 180 were the dominant congeners in the liver samples of the analysed animals, whereas PCB 138 and 153 in the lungs. TEQs levels calculated for only dioxin-like PCBs were low: 0.32 and 0.29 pg WHO-PCB-TEQ·g^?1 fat in liver of red deer and roe deer, respectively. The mean PCB concentrations obtained in our study for organs of roe deer and red deer were several times lower than those reported elsewhere. These findings show that the investigated roe and red deer originated from an area with low levels of PCB contamination.     

Variation in offspring sex ratio among individual Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>) females of different quality

The Trivers–Willard model predicts that in polygynous species, superior-quality females will maximize their fitness by producing male offspring. Using a sample of 1,780 Weddell seal (Leptonychotes weddellii) pups recorded over 31 years, we investigated relationships between offspring sex ratio and maternal age, reproductive experience, an index of maternal lifetime reproductive output, and annual environmental variations. We found evidence that females with higher index of lifetime reproductive output were more likely to produce male than female offspring but found only weak evidence that large-scale environmental variations influenced sex ratios. Our results suggest that mothers manipulate offspring sex to maximize their own fitness, and inherent maternal quality may influence offspring sex. These findings support the Trivers–Willard sex-allocation model. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Use of sibling relationship reconstruction to complement traditional monitoring in fisheries management and conservation of brown trout

Declining trends in the abundance of many fish urgently call for more efficient and informative monitoring methods that would provide necessary demographic data for the evaluation of existing conservation, restoration, and management actions. We investigated how genetic sibship reconstruction from young‐of‐the‐year brown trout (Salmo trutta L.) juveniles provides valuable, complementary demographic information that allowed us to disentangle the effects of habitat quality and number of breeders on juvenile density. We studied restored (n = 15) and control (n = 15) spawning and nursery habitats in 16 brown trout rivers and streams over 2 consecutive years to evaluate the effectiveness of habitat restoration activities. Similar juvenile densities both in restored and control spawning and nursery grounds were observed. Similarly, no differences in the effective number of breeders, N_b(SA), were detected between habitats, indicating that brown trout readily used recently restored spawning grounds. Only a weak relationship between the N_b(SA) and juvenile density was observed, suggesting that multiple factors affect juvenile abundance. In some areas, very low estimates of N_b(SA) were found at sites with high juvenile density, indicating that a small number of breeders can produce a high number of progeny in favorable conditions. In other sites, high N_b(SA) estimates were associated with low juvenile density, suggesting low habitat quality or lack of suitable spawning substrate in relation to available breeders. Based on these results, we recommend the incorporation of genetic sibship reconstruction to ongoing and future fish evaluation and monitoring programs to gain novel insights into local demographic and evolutionary processes relevant for fisheries management, habitat restoration, and conservation.     

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.     

Modelling landscape effects on density-contact rate relationships of deer in eastern Alberta: Implications for chronic wasting disease

Managing wildlife diseases requires an understanding of disease transmission, which may be strongly affected by host population density and landscape features. Transmission models are typically fit from time-series disease prevalence data and modelled based on how the contact rate among hosts is affected by density, which is often assumed to be a linear (density-dependent transmission) or constant (frequency-dependent transmission) relationship. However, long-term time-series data is unavailable for emerging diseases, and this approach cannot account for independent effects of landscape. We developed a mechanistic model based on ecological data to empirically derive the contact rate-density relationship in white-tailed and mule deer in an enzootic region of chronic wasting disease (CWD) in Alberta, Canada and to determine whether it was affected by landscape. Using data collected from aerial surveys and GPS-telemetry, we developed empirical relationships predicting deer group size, home range size, and habitat selection to iteratively simulate deer distributions across a range of densities and landscapes. We calculated a relative measure of total per-capita contact rate, which is proportional to the number of other deer contacted per individual per unit time, for each distribution as the sum of pairwise contact rates between a target deer and all other individuals. Each pairwise contact rate was estimated from an empirical relationship developed from GPS-telemetry data predicting pairwise contact rates as a function of home range overlap and landscape structure. Total per-capita contact rates increased as a saturating function of density, supporting a transmission model intermediate between density- and frequency-dependent transmission. This pattern resulted from group sizes that reached an asymptote with increasing deer density, although this relationship was mediated by tree and shrub coverage in the landscape, such that in heavily wooded areas, the contact rate saturated at much lower densities. These results suggest that CWD management based on herd reductions, which require a density-dependent contact rate to be effective, may have variable effects on disease across a single management region. The novel mechanistic approach we employed for estimating effects of density and landscape on transmission is a powerful complement to typical data-fitting approaches for modelling disease transmission.     

Contradictory findings in studies of sex ratio variation in roe deer (Capreolus capreolus)

Patterns of sex ratio variation and maternal investment reported in the literature are often inconsistent. This could be due to intra- and inter-specific variation in social systems, but may also be a result of the a posteriori nature of much of this type of analysis or the testing of models which are inappropriate. Two recent papers reported directly opposed results concerning variation in offspring sex ratios in relation to maternal condition in roe deer, interpreting the results as support for the Trivers and Willard model and for the local resource competition hypothesis, respectively. In this paper, we present data on offspring sex ratios and early juvenile body weight from two long-term studies of this species to test predictions arising from these two models concerning sex biases in litter composition and maternal care. First, we observed no consistent pattern of sex differences in an index of weaning weight or body weight at 1 month old in either population, indicating a lack of sex bias in maternal care. However, in one population, higher maternal body weight was associated with higher juvenile body weight of daughters, but not of sons. Secondly, we found a negative, but not statistically significant, relationship between maternal body weight and litter sex ratio such that heavier females tended to produce more daughters and lighter females to produce more sons. These results indicate that roe females which have additional investment potential available do not invest it in sons, as predicted by the Trivers and Willard model. Our results may provide some support that roe deer are subject to local resource competition acting at the level of the individual mother; however, the fact that particular trends in sex ratio data can be explained in functional terms provides no indication that they are actually adaptive. Received: 9 December 1997 / Accepted after revision: 11 November 1998     

Nest-site selection of passerines: effects of geographic scale and public and personal information

Nest-site selection is an important determinant of individual fitness in birds. Understanding what information individuals use to choose nest sites is therefore important for understanding the evolution of nest-site selection, the dynamics of populations, and the conservation of species. We used five years of mark-recapture data for Mountain Bluebirds (Sialia currucoides) to examine how dispersal probability and nest-site selection vary with potential cues of nest-site quality. Dispersal distance between breeding seasons and nest-site selection were modeled as a function of personal reproductive success, conspecific density, conspecific reproductive success, and habitat type. Between years, the dispersal probability was related to personal reproductive success, not conspecific information, and individuals fledging fewer young dispersed longer distances. For dispersing individuals, the probability that a nest site was selected in year i was negatively related to distance from the nest site selected in year i - 1 for all age and sex classes, and positively related to conspecific density and reproductive success in year i - 1 for both second-year (SY) and after-hatch-year (AHY) females. However, nest-site selection in year i was more strongly related to conspecific density in year i- 1 for hatch-year (HY) females and was much more strongly related to the reproductive success of conspecifics in year i - 1 for AHY females. Nest-site selection of HY and AHY males was not consistently related to the metrics of conspecific information, but we suspect that relationships were obscured by competitive interactions. We found no evidence indicating that individuals respond differently to conspecific information at longer distances, suggesting that individuals limit dispersal to areas where they have prior knowledge. We predict that these patterns of nest-site selection will allow birds to loosely track nest-site quality and maintain an ideal free distribution, where average fitness is equal in all habitat types.     

Social rank, fecundity and lifetime reproductive success in wild European rabbits (Oryctolagus cuniculus)

Wild European rabbits of both sexes have separate linear rank orders, which are established and maintained by intensive fights. This paper presents data from a 14-year study (1987-2000) on the population physiology and behavioural ecology of wild rabbits living in a 22,000 m² enclosure and focuses on the relationship between social rank and reproduction in females. Group composition, social ranks, fecundity and reproductive success were known for all females (n=197) from the outset of their first reproductive season at an age of about 300 days until their death. The annual reproductive success of females was influenced to a large extent by their social rank. This depended on two effects of about equal strength: a higher fecundity of high-ranking females and a lower mortality of their offspring between birth and adulthood. The lifetime reproductive success of the females varied greatly. Only about 50% of all females had any reproductive success (range: on to nine adult offspring). The social rank achieved by the females during their first reproductive season substantially influenced their lifetime reproductive success: The mean reproductive lifespan and lifetime fitness of high-ranking females (ranks 1 and 2) was about 60% greater than that of lower-ranking females, although many of the latter also gained dominant positions in subsequent years.     

Reproductive Investment of a Lacertid Lizard in Fragmented Habitat

Abstract:  We studied the effect of habitat fragmentation on female reproductive investment in a widespread lacertid lizard (  Psammodromus algirus ) in a mixed-forest archipelago of deciduous and evergreen oak woods in northern Spain. We captured gravid females in fragments (≤10 ha) and forests (≥ 200 ha) and brought them to the laboratory, where they laid their eggs. We incubated the eggs and released the first cohort of juveniles into the wild to monitor their survival. Females from fragments produced a smaller clutch mass and laid fewer eggs (relative to mean egg mass) than females of similar body size from forests. Lizards did not trade larger clutches for larger offspring, however, because females from fragments did not lay larger eggs (relative to their number) than females from forests. Among the first cohort of juveniles, larger egg mass and body size increased the probability of recapture the next year. Thus, fragmentation decreased the relative fecundity of lizards without increasing the quality of their offspring. Reduced energy availability, increased predation risk, and demographic stochasticity could decrease the fitness of lizards in fragmented habitats, which could contribute to the regional scarcity of this species in agricultural areas sprinkled with small patches of otherwise suitable forest. Our results show that predictable reduction of reproductive output with decreasing size of habitat patches can be added to the already known processes that cause inverse density dependence at low population numbers.     

Roe and red deer as bioindicators of heavy metals contamination in north-western Poland

The present paper examines the degree of environmental contamination in areas covered by the Natura 2000 programme, located in north-western Poland, with selected heavy metals based on their concentration in target organs of roe and red deer. Lead, cadmium, copper, iron, and zinc concentrations were determined by inductively coupled plasma–atomic emission spectrometry. Residues of lead and cadmium were found in most of the analysed samples of roe and red deer organs The concentration of cadmium in the organs of the animals studied was much higher than that of lead. The median for Pb in liver and kidneys was 0.055 and 0.092 μg/g dry weight (d.w.) in roe deer, and 0.067 and 0.081 μg/g d.w. in red deer, respectively. The median for liver and kidney cadmium was 0.770 and 6.139 μg/g d.w. in roe deer, and 0.422 and 6.365 μg/g d.w. in red deer, respectively. Our study has demonstrated that this area is laden with lead and cadmium. This is evidenced by the fact that maximum permissible levels of these elements in the organs of red and roe deer, which were used as bioindicators of environmental contamination, were exceeded.     

Socially induced delayed primiparity in brown bears Ursus arctos

Reproductive suppression through behavioral or physiological means is common in group-living and cooperative breeding mammals, but to our knowledge it has not been shown in wild large carnivores other than those with a clear form of social organization. Brown bear (Ursus arctos) females form matrilinear assemblages with related females using a common and largely exclusive area. Behavioral reproductive suppression might develop due to a hierarchical system among females within a matrilinear assemblage or due to inbreeding avoidance, because male brown bears can overlap with their daughters. We tested whether natal dispersal influenced the age of primiparity. We predicted that emigrant females, geographically removed from maternal or paternal influence, would reproduce earlier than philopatric females. The average age of primiparity was 4.3 years in females that dispersed outside their mother’s home range (n=8) and 5.2 years in philopatric females (n=10). Only the overlap with the mother’s home range, and not body size, body mass, growth, local population density, or overlap with the father’s home range, had a significant influence on the age of primiparity. The ultimate role of reproductive suppression for brown bears is likely to avoid inbreeding or to minimize resource competition. Due to the low risk of inbreeding and frequent exposure of young females to unrelated males, we conclude that resource competition within female hierarchies causes reproductive suppression in young females.     

Simulating Retention of Rare Alleles in Small Populations to Assess Management Options for Species with Different Life Histories

Preserving allelic diversity is important because it provides the capacity for adaptation and thus enables long‐term population viability. Allele retention is difficult to predict in animals with overlapping generations, so we used a new computer model to simulate retention of rare alleles in small populations of 3 species with contrasting life‐history traits: North Island Brown Kiwi (Apteryx mantelli; monogamous, long‐lived), North Island Robins (Petroica longipes; monogamous, short‐lived), and red deer (Cervus elaphus; polygynous, moderate lifespan). We simulated closed populations under various demographic scenarios and assessed the amounts of artificial immigration needed to achieve a goal of retaining 90% of selectively neutral rare alleles (frequency in the source population = 0.05) after 10 generations. The number of immigrants per generation required to meet the genetic goal ranged from 11 to 30, and there were key similarities and differences among species. None of the species met the genetic goal without immigration, and red deer lost the most allelic diversity due to reproductive skew among polygynous males. However, red deer required only a moderate rate of immigration relative to the other species to meet the genetic goal because nonterritorial breeders had a high turnover. Conversely, North Island Brown Kiwi needed the most immigration because the long lifespan of locally produced territorial breeders prevented a large proportion of immigrants from recruiting. In all species, the amount of immigration needed generally decreased with an increase in carrying capacity, survival, or reproductive output and increased as individual variation in reproductive success increased, indicating the importance of accurately quantifying these parameters to predict the effects of management. Overall, retaining rare alleles in a small, isolated population requires substantial investment of management effort. Use of simulations to explore strategies optimized for the populations in question will help maximize the value of this effort. Simulación de la Retención de Alelos Raros en Poblaciones Pequeñas para Evaluar Opciones de Manejo para Especies con Historias de Vida Diferentes     

A Natural Experiment on the Impact of Overabundant Deer on Forest Invertebrates

Abstract:  In large parts of North America and Europe, deer overabundance threatens forest plant diversity. Few researchers have examined its effects on invertebrate assemblages. In a natural experiment on Haida Gwaii (British Columbia, Canada), where Sitka black-tailed deer ( Odocoileus hemionus sitkensis ) were introduced, we compared islands with no deer, with deer for fewer than 20 years, and with deer for more than 50 years. We sampled invertebrates in three habitat categories: forest edge vegetation below the browse line, forest interior vegetation below the browse line, and forest interior litter. In forest edge vegetation, invertebrate abundance and species density decreased with increasing length of browsing history. In forest interior vegetation, decrease was significant only on islands with more than 50 years of browsing. Insect abundance in the vegetation decreased eightfold and species density sixfold on islands browsed for more than 50 years compared with islands without deer. Primary consumers were most affected. Invertebrates from the litter showed little or no variation related to browsing history. We attributed the difference between vegetation-dwelling and litter-dwelling invertebrates to differences in the effect of browsing on their habitat. In the layer below the browse line deer progressively removed the habitat. The extent of litter habitat was not affected, but its quality changed. We recommend more attention be given to the effect of overabundant ungulates on forest invertebrate conservation with a focus on edge and understory vegetation in addition to litter habitat.     

Nonideal breeding habitat selection: a mismatch between preference and fitness

The selection of breeding sites in heterogeneous habitats should ideally be based on cues closely reflecting habitat quality and thus predicting realized individual fitness. Using long-term population data and data on territory establishment of male Northern Wheatears (Oenanthe oenanthe), we examined whether territory characteristics linked to individual fitness (reproductive performance and survival) also were linked to territory preference. Breeding territories varied in their physical characteristics and their potential effects on reproductive performance, and this variation among territories was correlated from one year to the next. Of all measured territory characteristics (from the focal and the previous year) only territory field layer height predicted individual fitness, i.e., reproductive performance was higher in territories with permanently short rather than growing field layers. Territory preference, instead, was only linked to the size of territory aggregations, i.e., males settled earlier at territory sites sharing borders with several adjacent sites than at those with few or no adjacent sites. This mismatch between territory characteristics linked to fitness and those linked to territory preference was not explained by site fidelity or compensated for by the different fitness components measured. Because the results were not in agreement with an ecological trap scenario, where poor habitats are preferred over high-quality habitats, our results suggest a more general case of nonideal habitat selection. Whereas nonideal selection with respect to territory field layer height may be explained by its poor temporal predictability within the breeding season, the preference for territory aggregations is still open to alternative adaptive explanations. Our study suggests that nonideal habitat selection should be investigated by direct estimates of preferences (e.g., order of territory establishment) and their links to habitat characteristics and fitness components. Furthermore, we suggest that the probability of establishing a territory needs to be included as a factor influencing patterns of habitat selection.