Petter Kjellander

Higher levels of multiple ecosystem services are found in forests with more tree species

Forests are of major importance to human society, contributing several crucial ecosystem services. Biodiversity is suggested to positively influence multiple services but evidence from natural systems at scales relevant to management is scarce. Here, across a scale of 400,000 km2, we report that tree species richness in production forests shows positive to positively hump-shaped relationships with multiple ecosystem services. These include production of tree biomass, soil carbon storage, berry production and game production potential. For example, biomass production was approximately 50% greater with five than with one tree species. In addition, we show positive relationships between tree species richness and proxies for other biodiversity components. Importantly, no single tree species was able to promote all services, and some services were negatively correlated to each other. Management of production forests will therefore benefit from considering multiple tree species to sustain the full range of benefits that the society obtains from forests.

Variations in adult body mass in roe deer: the effects of population density at birth and of habitat quality

Body mass is a key determinant of fitness components in many organisms, and adult mass varies considerably among individuals within populations. These variations have several causes, involve temporal and spatial factors, and are not yet well understood. We use long‐term data from 20 roe deer cohorts (1977‐96) in a 2600 ha study area (Chizé, western France) with two habitats contrasting in quality (rich oak forest in the North versus poor beech forest in the South) to analyse the effects of both cohort and habitat quality on adult mass (i.e. median body mass between 4 and 10 years of age) of roe deer (Capreolus capreolus). Cohort strongly influenced the adult body mass of roe deer in both sexes: males born in 1994 were 5.2 kg heavier when aged between 4 and 10 years old than males born in 1986, while females born in 1995 were 4.7 kg heavier between 4 and 10 years old than females born in 1982. For a given cohort, adult males were, on average, 0.9 kg heavier in the rich oak forest than in the poor beech forest. A similar trend occurred for adult females (0.5 kg heavier in the oak forest). The effects of cohort and habitat were additive and accounted for ca. 40% of the variation observed in the adult mass of roe deer at Chizé (males: 41.2%; females: 40.2%). Population density during the spring of the birth accounted for about 35% of cohort variation, whereas rainfall in May‐June had no effect. Such delayed effects of density at birth on adult body mass probably affect population dynamics, and might constitute a mechanism by which delayed density‐dependence occurs in ungulate populations.

Antler size provides an honest signal of male phenotypic quality in roe deer.

Identifying factors shaping secondary sexual traits is essential in understanding how their variation may influence male fitness. Little information is available on the allocation of resources to antler growth in territorial ungulates with low sexual size dimorphism. We investigated phenotypic and environmental factors affecting both absolute and relative antler size of male roe deer in three contrasting populations in France and Sweden. In the three populations, we found marked age‐specific variation in antler size, with an increase in both absolute and relative antler size between yearling and prime‐age stages, followed by a decrease (senescence) for males older than 7 years. Antler size increased allometrically with body mass. This increase was particularly strong for senescent males, suggesting the evolution of two reproductive tactics: heavy old males invested particularly heavily in antler growth (potentially remaining competitive for territories), whereas light old males grew small antlers (potentially abandoning territory defense). Finally, environmental conditions had little effect on antler size: only population density negatively affected absolute antler size in one of the three populations. Antler size may therefore provide an honest signal of male phenotypic quality in roe deer. We discuss the implications of these results in terms of territory tenure and mating competition.

Ideal free distribution and natal dispersal in female roe deer

We investigated whether adult (≧ 2 years) female roe deer Capreolus capreolus conform to an ideal free or an ideal despotic distribution, in order to understand whether natal dispersal is ‘voluntary’ or socially enforced. The study was undertaken in a high-density, free-ranging population close to Stockholm, Sweden, during 1989–1994. Data on population density, habitat quality, and five parameters on female reproduction and body condition, in addition to age distribution, were obtained in two nearby located areas, the ‘field’ and the ‘forest’, representing contrasting habitats. Population density was estimated by faecal pellet group counting in addition to total counts of culled animals after a major deer harvest. Density in the ‘field area’ was twice that in the ‘forest area’ (66 vs 33 deer/km2). Habitat quality was determined by analysing the amount of faecal nitrogen from samples of faeces collected in the two areas, and by comparing size of summer home ranges for adults of both sexes with the aid of telemetry. Both estimates indicated a higher nutritional quality and resource abundance, respectively, in the ‘field area’, with faecal nitrogen content being higher (2.96 vs 2.43%), and ranges being smaller (12.9 vs 20.9 ha). No significant differences were found in any of the parameters on reproduction or body condition obtained from culled females, i.e. number of corpora lutea (1.8 vs 1.8), proportion of females with offspring (71 vs 56%), body mass (16.9 vs 17.3 kg), kidney fat index (91.9 vs 98.5), and length of the lower mandible (152.8 vs 151.4 mm). Neither did the age distribution among females, as determined from tooth wear, differ between the areas. These results are consistent with the prediction of the ideal free hypothesis. This, in turn, suggests that female dispersal in this species is ‘voluntary’, the underlying proximate cause being maximization of resource gain.

Personality and foraging decisions in fallow deer, Dama dama

Recent studies have examined the ecological and evolutionary bases for variation in animal personality. However, only a few such studies have examined how foraging parameters are influenced by different personality domains. In wild ungulates, the trade-off between the time spent on food intake and antipredator behaviour differs between individuals, but the underlying reason for this is not yet well understood. One possibility is that this trade-off reflects personality dimensions such as boldness. To relate foraging decisions to personality we measured personality and performed feeding experiments with familiar and novel food in familiar and novel situations. We measured personality traits in 15 tame fallow deer, using novel object tests (NO), behavioural observations (BO) and personality ratings (PR). Boldness dimensions were found using PR and NO, dominance dimensions were found using BO and PR, and a flexibility dimension was found using BO. Multitrait–multimethod analysis showed that similar dimensions were significantly correlated across different methods and that different dimensions were not significantly correlated, even if measured using the same method. We also found that novel food eaten in familiar situations and familiar food eaten in novel situations were strongly related to boldness but not dominance, flexibility or age. Thus the trade-off between the benefits of gaining more food and the costs of reduced vigilance or increased toxin ingestion reflect boldness. These findings highlight the nature of personality dimensions in ungulates and how boldness impacts foraging behaviour.

Quantitative estimates of tree species selectivity by moose (Alces alces) in a forest landscape

Abstract An extensive literature is available on browsing preference for certain tree species. However, useful predictive tools for estimating the impact of deer on forests production and biodiversity can still be improved. A step in that direction is not only to rank preference among tree species but also to quantify the relative risk of being browsed. The foraging selectivity of moose was evaluated using three different statistical methods developed to study habitat utilization. The general pattern for the three methods was consistent. From the results, groups of forage species were clustered and a quantitative index of selectivity was calculated for the groups. The selectivity index showed that rowan (Sorbus aucuparia), willow (Salix ssp.) and aspen (Populus tremula) had a 14 times higher probability of being browsed than a group consisting of Scots pine (Pinus sylvestris) and downy birch (Betula pubescens), while juniper (Juniperus communis) and silver birch (Betula pendula) had a 3.5 times higher probability than Scots pine and downy birch. Since the most preferred species were the least abundant, one should be cautious about the generality of the index between areas, as it may indicate that preference depends on plant species composition. The method used can easily be applied in forest management. Information on quantitative selectivity indices may improve the possibility of managing moose in accordance with acceptable browsing damage.

Archiving primary data: solutions for long-term studies

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

Seasonality, weather and climate affect home range size in roe deer across a wide latitudinal gradient within Europe

1. Because many large mammal species have wide geographical ranges, spatially distant populations may be confronted with different sets of environmental conditions. Investigating how home range (HR) size varies across environmental gradients should yield a better understanding of the factors affecting large mammal ecology. 2. We evaluated how HR size of a large herbivore, the roe deer (Capreolus capreolus), varies in relation to seasonality, latitude (climate), weather, plant productivity and landscape features across its geographical range in Western Europe. As roe deer are income breeders, expected to adjust HR size continuously to temporal variation in food resources and energetic requirements, our baseline prediction was for HR size to decrease with proxies of resource availability. 3. We used GPS locations of roe deer collected from seven study sites (EURODEER collaborative project) to estimate fixed-kernel HR size at weekly and monthly temporal scales. We performed an unusually comprehensive analysis of variation in HR size among and within populations over time across the geographical range of a single species using generalized additive mixed models and linear mixed models, respectively. 4. Among populations, HR size decreased with increasing values for proxies of forage abundance, but increased with increases in seasonality, stochastic variation of temperature, latitude and snow cover. Within populations, roe deer HR size varied over time in relation to seasonality and proxies of forage abundance in a consistent way across the seven populations. Thus, our findings were broadly consistent across the distributional range of this species, demonstrating a strong and ubiquitous link between the amplitude and timing of environmental seasonality and HR size at the continental scale. 5. Overall, the variability in average HR size of roe deer across Europe reflects the interaction among local weather, climate and seasonality, providing valuable insight into the limiting factors affecting this large herbivore under contrasting conditions. The complexity of the relationships suggests that predicting ranging behaviour of large herbivores in relation to current and future climate change will require detailed knowledge not only about predicted increases in temperature, but also how this interacts with factors such as day length and climate predictability.

Population density and sex do not influence fine-scale natal dispersal in roe deer.

It is commonly assumed that the propensity to disperse and the dispersal distance of mammals should increase with increasing density and be greater among males than among females. However, most empirical evidence, especially on large mammals, has focused on highly polygynous and dimorphic species displaying female-defence mating tactics. We tested these predictions on roe deer, a weakly polygynous species of large herbivore exhibiting a resource-defence mating tactic at a fine spatial scale. Using three long-term studies of populations that were subject to the experimental manipulation of size, we did not find any support for either prediction, whether in terms of dispersal probability or dispersal distance. Our findings of similar dispersal patterns in both sexes of roe deer suggest that the underlying cause of natal dispersal is not related to inbreeding avoidance in this species. The absence of positive density dependence in fine-scale dispersal behaviour suggests that roe deer natal dispersal is a pre-saturation process that is shaped by heterogeneities in habitat quality rather than by density per se.

AGE-SPECIFIC VARIATION IN MALE BREEDING SUCCESS OF A TERRITORIAL UNGULATE SPECIES, THE EUROPEAN ROE DEER

Abstract We investigated age-specific variation in male yearly breeding success (YBS) using genetic estimates obtained from 2 populations of a territorial ungulate, the European roe deer (Capreolus capreolus). YBS in both populations was markedly age-structured, with 3 distinct stages, supporting the dome-shaped pattern of variation commonly reported for age-dependent variation in life-history traits of ungulates. YBS was low at 2 years of age, peaked at 3–8 years of age, and tended to decline afterwards (senescence). Most males successfully reproduced for the 1st time at 3 years of age, which is well after their physiological maturity. The few successful young males (i.e., 2 year olds) were likely fast-growing individuals that could successfully hold a territory. The high variance in YBS and antler size for old males at Bogesund, Sweden, suggests that only some males of this age class are able to maintain large antlers and, hence, retain their territories.