Elevated CO2 alters deployment of roots in “small” growth containers

Since European badgers (Meles meles L.) form non-cooperative groups in parts of their geographic range, but are solitary elsewhere, their social systems have been at the centre of a debate about the evolution of group living in the Carnivora. In a recent review of models of non-cooperative sociality, Woodroffe and Macdonald (1993) presented evidence in favour of two hypotheses, which suggested that badger groups might form because either the distribution of blocks of foodrich habitat, or the economics of excavating new setts, prevented the division of group territories into individual territories. We present data upon the response of badger spatial organisation to a reduction in food-patch dispersion, brought about by the conversion of carthwormpoor arable land to earthworm-rich pasture over a 15-year period. This change in the distribution of earthworm-rich habitats was accompanied by territory fission, facilitated by the excavation of new setts. This indicates that the availability of sett sites had not constrained territory size at the start of the study. However, sett distribution did define the size and configuration of the daughter territories. We also show that variation among territories in the availability of food-rich habitats was reflected in the reproductive rates and body weights of the groups that inhabited them, although there was no detectable effect upon group size.     

Using the Mahalanobis distance statistic with unplanned presence-only survey data for biogeographical models of species distribution and abundance: a case study of badger setts

Aim Project‐specific data for biogeographical models are often logistically impractical to collect, forcing the use of existing data from a variety of sources. Use of these data is complicated when neither absence nor an estimate of the area sampled is available, as these are requirements of most analytical techniques. We demonstrate the Mahalanobis distance statistic (D²), which is a presence‐only modelling technique and does not require information on species absence or the sampled area. We use badger (Meles meles) setts as the basis for this investigation, as their landscape associations are well understood, and survey data exist against which to compare estimates of sett distribution and abundance. Location  England and Wales (151,403 km²). Methods We used stratified random samples of sett locations, and landscape variables that are known to be important for choice of badger sett location within a geographic information system at a cell resolution of 100 × 100 m. Landscape conditions at two scales were extracted, at and around sett locations, and the D² was used to classify all cells in England and Wales into a sett suitability model. Comparison of this sett suitability model with known main sett densities allowed estimates of main sett density to be made across England and Wales, with associated uncertainty. Results The sett suitability model was shown through iterative sampling and model evaluation using independent data to be stable and accurate. Main sett density estimates were biologically plausible in comparison with previous field‐derived estimates. We estimate 58,000 main setts within England and Wales, with 95% confidence intervals suggesting a value between 31,000 and 93,000. Main conclusions The D², which could be applied to other species and locations, proved useful in our context, where absence data were not available and the sampled area could not be reliably established. We were able to predict sett suitability across a large area and at a fine resolution, and to generate plausible estimates of main sett density. The final model provides valuable information on probable badger sett distribution and abundance, and may contribute to future research on the spatial ecology of badgers in England and Wales.     

Seasonal and spatial pattern of shelter use by badgers<Emphasis Type="Italic">Meles meles</Emphasis> in Białowieża Primeval Forest (Poland)

In 1997–2001, we investigated the use of day-time shelters by radio-collared badgersMeles meles (Linnaeus, 1758) in the Białowieża Primeval Forest, eastern Poland. Each social group of badgers utilised, on average, 9 different shelters per territory (range: 4–20). The main setts, occupied for breeding and winter sleep, were also most frequently used for day-time rest throughout the year (73% of days). Badgers living in the pristine oldgrowth stands utilised larger number of shelters and spent more days in hollow trees (mainly limeTilia cordata), compared to badgers inhabiting younger secondary tree stands. Number of shelters used by individuals varied between seasons and depended on sex and age of animals. In summer, badgers used more shelters than in spring and autumn. In winter, they stayed in their main setts only. Adult males occupied more shelters and spent fewer days in the main sett than other badgers. In spring, females rearing young used only the main setts. The average underground space used by badgers within the main sett was 128 m². It was largest in summer and smallest in winter, and also varied between males and females. We proposed that, in a low-density population, badgers used several setts and other daily shelters to reduce energy expenditure when exploring their large territories and foraging. Furthermore, setts may play a role of marking sites. Analysis of the biogeographical pattern of sett use by European badgers showed that the number of setts used by social groups increased with increasing territory size, whereas the density of setts (n setts/km²) was negatively correlated with territory size. We proposed that different factors could shape the utilisation of setts by badgers in low- and high-density populations.     

Internal structure and contents of three badger (Meles meles) setts

Three badger ( Meles meles L.) setts in the south of England, which formed a single sett complex belonging to one social group of badgers, were excavated prior to being destroyed by construction of a new road. Setts 1 and 2, classified as annexes, were excavated completely; sett 3, classified as a main sett, was only excavated partially, but its total size was estimated from the excavated portion. The setts consisted of tunnels totalling 16 m, 140 m and 879 m, respectively; contained one, nine and 50 chambers; and had five, 42 and 178 entrances. The total volume of the three setts was about 45 m³, and their construction was estimated to have required the removal of about 70 tonnes of soil. In the two smaller setts tunnels ran on a single level with an average depth of 99 cm; in the larger sett they ran on two levels with modal depths of 50 cm and 110–120 cm, respectively. All three setts contained bedding material (dry grass and plastic bags) but only the main sett contained latrines. None of the setts contained badger bones and the interiors of all three setts were remarkably clean and orderly. We discuss hypotheses as to why badgers sometimes continue to extend even large well-established setts but conclude that the survival value of very large setts remains problematical.     

Predicting Distribution and Density of European Badger (<Emphasis Type="Italic">Meles Meles</Emphasis>) Setts in Denmark

Predictive models of the spatial distribution and abundance of species based on habitat characteristics are finding increasing use in management and conservation. The European badger attracts interest as a model species both for conservation reasons and because of the important role the species is playing in understanding carnivore sociality. We developed a statistical habitat model based on presence/absence data on badger setts. To maximise the utility of the model in management, we limited the choice of model variables to those that had a clear basis in badger ecology and that could be obtained on a nation-wide digital format. We extrapolated the habitat model to a region in Denmark and developed a threshold-independent sett distribution algorithm to estimate sett densities. The habitat model was simpler than previously published models of badger sett habitat selection, but nevertheless had a predictive ability in excess of 80% judged against independent data. The sett distribution algorithm was able to simultaneously reproduce several observed patterns of sett density and distribution over the probability gradient. It thus represents a significant improvement over threshold-dependent methods used to discriminate between suitable and unsuitable habitat predicted by presence/absence regression models. Our approach demonstrates that a model of badger sett habitat suitability with high predictive power can be obtained using easily accessible map-variables and presence/absence data. This is a prerequisite for using habitat models as predictive tools over large areas. The use of a simple sett distribution algorithm circumvents the common problem of subjectively fixing a threshold to discriminate between suitable and unsuitable habitat. In conjunction the models presented here constitute an important contribution to the management of the badger in Denmark and, upon further validation, possibly to similar regions in Northern Europe.     

Thiessen polygons as a model for animal territory estimation

Thiessen polygons are often used to model territory characteristics. However, information about the quality of Thiessen polygon‐based estimates is currently lacking. We used published data to investigate the match between Thiessen polygons and mapped bird territories regarding territory size, shape and neighbourhood. Although territory sizes and the number of neighbours were strongly correlated between these two methods, both parameters were overestimated by the Thiessen polygons. Therefore, caution is required when Thiessen polygons are used as a model for absolute values and when the assumptions of Thiessen polygons, such as formation of discrete territories and a contiguous study area, are not met.     

Field evaluation of abundance estimates under binomial and multinomial N-mixture models

Assessing and modelling abundance from animal count data is a very common task in ecology and management. Detection is arguably never perfect, but modern hierarchical models can incorporate detection probability and yield abundance estimates that are corrected for imperfect detection. Two variants of these models rely on counts of unmarked individuals, or territories (binomial N-mixture models, or binmix), and on detection histories based on territory-mapping data (multinomial N-mixture models or multimix). However, calibration studies which evaluate these two N-mixture model approaches are needed. We analysed conventional territory-mapping data (three surveys in 2014 and four in 2015) using both binmix and multimix models to estimate abundance for two common avian cavity-nesting forest species (Great Tit Parus major and Eurasian Blue Tit Cyanistes caeruleus). In the same study area, we used two benchmarks: occupancy data from a dense nestbox scheme and total number of detected territories. To investigate variance in estimates due to the territory assignment, three independent ornithologists conducted territory assignments. Nestbox occupancy yields a minimum number of territories, as some natural cavities may have been used, and binmix model estimates were generally higher than this benchmark. Estimates using the multimix model were slightly more precise than binmix model estimates. Depending on the person assigning the territories, the multimix model estimates became quite different, either overestimating or underestimating the ‘truth’. We conclude that N-mixture models estimated abundance reliably, even for our very small sample sizes. Territory-mapping counts depended on territory assignment and this carried over to estimates under the multimix model. This limitation has to be taken into account when abundance estimates are compared between sites or years. Whenever possible, accounting for such hidden heterogeneity in the raw data of bird surveys, via including a ‘territory editor’ factor, is recommended. Distributing the surveys randomly (in time and space) to editors may also alleviate this problem.     

Group size versus territory size in group-living badgers: a large-sample field test of the Resource Dispersion Hypothesis

Badgers ( Meles meles ) have been the focus for the development of a pervasive model of social grouping behaviour, relevant to a number of carnivore species and other taxonomic groups – the Resource Dispersion Hypothesis (RDH). The RDH hypothesises that the dispersion and richness of resources in the environment provide a passive mechanism for the formation of groups, even without any direct benefits of group living. However, few studies have tested the RDH in the field. The principal prediction is that, as opposed to enlargement of territory sizes to accommodate more members, territory size (TS) is independent of group size (GS). Instead, TS is determined by the spatial dispersion of resources, while GS is independently determined by the richness of those resources. However, these predictions provide only weak correlative tests, especially in non-experimental field studies. The first predicts an absence of correlation and is therefore prone to Type II error, especially given the small sample sizes and errors in estimating TS and GS of mammals in the field. We tested for independence of territory size and group size in all years with available data since the beginning of the long-term badger study in Wytham Woods in 1974. We used two methods of TS estimation, a sequential Bonferroni technique to adjust for multiple inference tests, a combined analysis and an analysis with pooled data. This prediction of the RDH could not be rejected on the basis of any of these analyses. Given this evidence that other processes are independently determining group size and territory size, further predictions of the RDH will be worth investigating in considerable detail.     

Mating systems and territory in Lapwings Vanellus vanellus

In a population of Lapwings Vanellus vanellus studied for 5 years near Bergen, southwester Norway, 23–41% of the males were polygynous, having two and sometimes three mates, whereas 50–77% were monogamous and 0–21% remained unmated. Bachelors held territories in or immediately adjacent to the study area each of the years. As predicted from the Polygyny Threshold model, primary females generally laid eggs earlier than monogamous females, but the overlap in laying dates was substantial between these two classes. The number of females and start of breeding on the territories were analysed in relation to nine variables pertaining to safety from predation and to food. Of these, territory size correlated most consistently with number of females (positively) and the laying of first egg (negatively). Numbers of females observed were not significantly different from numbers expected from territory size in 4 out of 5 years; however, earlier start of breeding on large territories indicated that females did not settle according to the Neutral Mate Choice model. The Female-biased Sex Ratio hypothesis was refuted, unmated territorial males being available throughout the breeding seasons.     

Drivers of sett site location by European badgers in Portugal

European badgers (Meles meles) are considered central-place foragers, whose spatial ecology is predominantly determined by sett location. Many studies have assessed the factors determining sett site selection throughout this species’ range, but these have often been geographically limited and have primarily identified locally dependent factors. To infer key factors determining sett location, a broader scale approach is needed. Between June 2014 and January 2017, we surveyed mainland Portugal to detect badger setts in 10?×?10 km cells, corresponding to a total of 657.5 km walked line transects. We detected 54 main setts in 136 surveyed cells. Each sett and non-sett site (i.e. transects without setts) was characterised using bioenvironmental variables (e.g. land cover, presence of human infrastructure, soil). We used generalized linear mixed models to test five hypotheses potentially explaining sett location: land cover composition; anthropogenic disturbance; abiotic environmental drivers; trophic resource availability; and a combined effect of all these factors. Our findings show that the key factors for badger sett site selection in Portugal are: (1) disturbance avoidance (low beehive density; absence of livestock; far from hunting reserves), but with a tendency to be located close to highways and unpaved roads; and (2) ease of excavation (avoidance of sedimentary/metamorphic composite rocks). Although specific factors among these drivers may be more important locally or regionally, these major drivers have also been identified elsewhere in Europe. Our nationwide approach contributes to a broader understanding of general patterns of sett site selection by badgers in southern Europe. Furthermore, it provides the national authorities with novel and broad-scale data to facilitate sustainable species conservation of badgers in the southwestern limit of their range.     

Escape Tactics and Alarm Responses in Badgers Meles meles: A Field Experiment

A typical badger (Meles meles) territory contains a primary burrow or 'main sett' plus several secondary burrows or 'outlier setts'. The main aim of our study was to test the hypothesis that outlier setts are used as emergency refuges, by subjecting foraging radio-collared badgers to three levels of experimental disturbance (low, moderate and high). In addition, we recorded the occurrence of potential alarm signals. With low-level disturbance, badgers usually returned to the main sett even when this was further away than the nearest outlier sett; with high-level disturbance they always took refuge in the nearest outlier; and with moderate-level disturbance they showed no clear preference. We conclude that outlier setts do act as emergency refuges, but only when an animal is badly frightened. Possible alarm signals (pilo-erection, head-flagging, snorting and growling) occurred mainly during moderate or high-level disturbance but even then they were relatively infrequent. Signals were no more likely to be emitted when conspecifics were near by than when the signaller was alone, and when conspecifics were present they rarely reacted either to the flight of the disturbed animal or to any signals that it emitted. We conclude that such alarm signals as do occur constitute threats directed towards the predator rather than warnings for the benefit of conspecifics.     

Comparison of the predicted impact of a screwworm fly outbreak in Australia using a growth index model and a life-cycle model

Abstract. The spatial population dynamics of an Old World screwworm fly, Chrysomya bezziana Villeneuve (OWS), outbreak in Australia have been modelled in two ways. The first model uses weekly growth indices derived from climatic data to predict the adult female population. The second is a detailed cohort life-cycle model. Due to technical and time constraints, the growth index model is preferred as the biological component of a much larger bioeconomic model because of its smaller program size and faster execution. In deciding whether adoption of the growth index model would be at the expense of scientific accuracy, the life-cycle model was developed as a yardstick. We showed that the growth index model was a practical and adequate substitution for the OWS life-cycle model and a novel spatial/temporal modelling approach with generic qualities. We elaborate on the previously reported growth index model, describe the life-cycle model and compare the results of both models. In the event of an OWS incursion in northern or eastern Australia, given average climatic conditions, both models predict that most of the suitable range (some 2.3M km²) will be colonized within 4–5 years if an eradication campaign is not attempted. Much of its permanent range would be in tropical and subtropical extensive grazing regions. Where computer or funding resources are restrictive, models incorporating growth indices may prove adequate for spatial population studies of some species.     

Allometric models for non-destructive leaf area estimation in coffee (Coffea arabica and Coffea canephora)

We aimed to evaluate the currently used allometric models, as well as to propose a reliable and accurate model using non-destructive measurements of leaf length (L) and/or width (W), for estimating the area of leaves of eight field-grown coffee cultivars. For model construction, a total of 1563 leaves were randomly selected from different levels of the tree canopies and encompassed the full spectrum of measurable leaf sizes (0.3–263 cm²) for each genotype. Power models better fit coffee leaf area (LA) than linear models. To validate the model, an independent data set of 388 leaves was used. We demonstrated that the currently used allometric models are biased, underestimating the area of a coffee leaf. We developed a single power model     based on two leaf dimensions [LA = 0.6626 (LW)^1.0116; standard errors: β₀ = 0.0064, β₁ = 0.0019; R² = 0.996] with high precision and accuracy, random dispersion pattern of residuals and also unbiased, irrespective of cultivar and leaf size and shape. Even when the L (but not width) alone was used as the single leaf dimension, the power model developed still predicted with good accuracy the LA but at the expense of some loss of precision, as particularly found for 8% of the leaves sampled with length-to-width ratios below 2.0 or above 3.0.     

Species distribution models as a tool to estimate reproductive parameters: a case study with a passerine bird species

1.?Correlative species distribution models (SDMs) assess relationships between species distribution data and environmental features, to evaluate the environmental suitability (ES) of a given area for a species, by providing a measure of the probability of presence. If the output of SDMs represents the relationships between habitat features and species performance well, SDM results can be related also to other key parameters of populations, including reproductive parameters. To test this hypothesis, we evaluated whether SDM results can be used as a proxy of reproductive parameters (breeding output, territory size) in red-backed shrikes (Lanius collurio). 2.?The distribution of 726 shrike territories in Northern Italy was obtained through multiple focused surveys; for a subset of pairs, we also measured territory area and number of fledged juveniles. We used Maximum Entropy modelling to build a SDM on the basis of territory distribution. We used generalized least squares and spatial generalized mixed models to relate territory size and number of fledged juveniles to SDM suitability, while controlling for spatial autocorrelation. 3.?Species distribution models predicted shrike distribution very well. Territory size was negatively related to suitability estimated through SDM, while the number of fledglings significantly increased with the suitability of the territory. This was true also when SDM was built using only spatially and temporally independent data. 4.?Results show a clear relationship between ES estimated through presence-only SDMs and two key parameters related to species' reproduction, suggesting that suitability estimated by SDM, and habitat quality determining reproduction parameters in our model system, are correlated. Our study shows the potential use of SDMs to infer important fitness parameters; this information can have great importance in management and conservation.     

Dynamic interactions among badgers: implications for sociality and disease transmission

1. Direct interactions between individuals play an important part in the sociality of group-living animals, their mating system and disease transmission. Here, we devise a methodology to quantify relative rates of proximity interaction from radio-tracking data and highlight potential asymmetries within the contact network of a moderate-density badger population in the north-east of England. 2. We analysed radio-tracking data from four contiguous social groups, collected over a 3-year period. Dynamic interaction analysis of badger dyads was used to assess the movement of individuals in relation to the movement of others, both within and between social groups. Dyads were assessed with regard to season, sex, age and sett use pattern of the badgers involved. 3. Intragroup separation distances were significantly shorter than intergroup separation distances, and interactions between groups were rare. Within groups, individuals interacted with each other more often than expected, and interaction patterns varied significantly with season and sett use pattern. Non-mover dyads (using the main sett for day-resting on > 50% of occasions) interacted more frequently than mover dyads (using an outlier sett for day-resting on > 50% of occasions) or mover-non-mover dyads. Interactions between group members occurred most frequently in winter. 4. Of close intragroup interactions (< 50 m separation distance), 88.6% were associated with a main sett and only 4.4% with outlier setts. Non-mover dyads and non-mover-mover dyads interacted significantly more often at the main sett than mover-only dyads. These results highlight the importance of the main sett to badger sociality and support the suggestion that badger social groups are comprised of different subgroups, in our case based on differential sett use patterns. 5. Asymmetries in contact structure within a population will affect the way in which diseases are transmitted through a social network. Assessment of these networks is essential for understanding the persistence and spread of disease within populations which do not mix freely or which exhibit heterogeneities in their spatial or social behaviour.     

Social Spacing in Small Mammals: Patterns of Individual Variation

The pattern of social spacing in small mammals differs fromthat observed in many other vertebrates. Small mammals frequentlyhave non-exclusive territories and tolerate a large amount ofoverlap with other conspecifics. The determinant factors ofhome range or territory size in small mammals are not knownfor most species. We carried out a study of the determinantfactors of home range size in a model small mammal, the easternchipmunk, Tamias striatus. The population was studied for fiveyears. The effect of experimental perturbations on food supplyand population density offered strong evidence that the meanhome range size in the population was determined by resourceabundance. Changes in population density had little or no measurableeffect. We noted that even when mean home range size decreasedsignificantly in response to an increase in available food,a great deal of variability in individual home range sizes remained.We hypothesized that this pattern of variation among individualswas also resource related; large home ranges would be locatedin areas of low resource density and small home ranges wouldbe located in areas of high resource density. Our data to datedo not offer support for this hypothesis; however our researchhas shown that the data needed to convincingly reject the nullhypothesis are very complex. We discuss the evidence requiredto study patterns of individual variation, and how models ofoptimal territory size may be useful. Research that examinespatterns of individual variation are few in number, yet studiesof individual variation will ultimately provide the best insightson the dynamics of evolutionary ecology.     

On the Bayesian estimation of a closed population size in the presence of heterogeneity and model uncertainty

Summary .   We consider the estimation of the size of a closed population, often of interest for wild animal populations, using a capture–recapture study. The estimate of the total population size can be very sensitive to the choice of model used to fit to the data. We consider a Bayesian approach, in which we consider all eight plausible models initially described by Otis et al. (1978, Wildlife Monographs 62, 1–135) within a single framework, including models containing an individual heterogeneity component. We show how we are able to obtain a model-averaged estimate of the total population, incorporating both parameter and model uncertainty. To illustrate the methodology we initially perform a simulation study and analyze two datasets where the population size is known, before considering a real example relating to a population of dolphins off northeast Scotland.     

Patterns of territory size and defence level in rural and urban tawny owl (Strix aluco) populations

The effect of different variables on the territory size and defence level of tawny owl ( Strix aluco L.) populations was studied in two habitats: mixed farmland (1976–1978) and town (1986–1988). Territory size was determined by plotting the position of territorial challenges (hoots) of males which were individually recognizable in most cases. No differences were found in size, defence, number of competitors and biomass between territories in farmland and in town, but territories in town were more fragmented and showed a larger nearest neighbour distance. Size of rural territories was adequately predicted both by some structural habitat features and food supplies, while size of urban territories was affected only by habitat structure and by years of territory occupancy. No effect on territory size by competing neighbours was found in either habitat, thus confirming that behavioural interactions between the owners of territories and potential settlers prevent a decrease of territory size.
Defence level of urban territories was affected by 'social'variables (nearest neighbour distance and years of occupancy), structural variables (percentage of poplar grove and diversity), and food supply, while in rural territories only structural variables (fragmentation or natural vegetation) and food supplies affected defence level.
This was due to different settlement tactics in the two areas, depending on the extent and quality of suitable habitat.     

Sett density as an estimator of population density in the European badger Meles meles

• 1 Estimations of European badger population density in the UK are usually based on surveys of numbers of main setts. However, this approach cannot be used in low‐density areas, such as the Mediterranean region, where no main setts can be defined. Therefore, an alternative method is needed to estimate badger density over large areas.
• 2 We reviewed the existing published information to evaluate whether badger density is correlated to the density of all setts in an area (not only the main setts) and to ask whether badger density can be predicted from total sett number throughout the geographical range of the species.
• 3 In multiple regression analysis, badger density and the size of the study area explained 73% of the variance in sett density. Badger density had a significant positive effect on sett density, while the effect of study area size was not significant. Therefore, total sett density can be used to obtain an estimation of relative badger density in all habitats and regions throughout the badger's geographical range, allowing comparative research.

     

Predation Risk,Resource Quality,and Reef Structural Complexity Shape Territoriality in a Coral Reef Herbivore

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual’s ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.