The potential of grass field margin management for enhancing beetle diversity in intensive livestock farms

1. Declining populations of UK grassland flora and fauna have been attributed to intensification of agricultural management practices, including changes in cutting, fertilizer, grazing and drainage regimes. We aimed to develop field margin management practices that could reverse declines in intensively managed grassland biodiversity that would have application in the UK and Europe. Here we focus on one aspect of grassland biodiversity, the beetles. 2. In four intensively managed livestock farms in south-west England, 10-m wide field margins in existing grasslands were managed to create seven treatments of increasing sward architectural complexity. This was achieved through combinations of inorganic (NPK) fertilizer, cattle grazing, and timing and height of cutting. To examine the potential influence of complexity on faunal diversity, beetles were identified to species level from suction samples taken between 2003 and 2005, and their assemblage structure was related to margin management, floral assemblages and sward architecture. 3. Beetle abundance, and species richness and evenness were influenced by margin management treatment and its interaction with year. Correlations with sward architecture and the percentage cover of dominant forbs and grasses were also found. Functional groups of the beetles showed different responses to the management treatments. In particular, higher proportional abundances of seed/flower-feeding guilds were found in treatments not receiving NPK fertilizer. 4. The assemblage structure was shown to respond to margin management treatments, sward architecture and the percentage cover of dominant forbs and grasses. The most extensively managed treatments were characterized by distinct successional trajectories from the control treatment. 5. Synthesis and applications. This study provides management options suitable for use within agri-environment schemes intended to improve faunal diversity associated with intensively managed lowland grasslands. Field margins receiving either no management or a single July silage cut were shown to support greater abundances and species richness of beetles, although subtler modifications of conventional management may also be beneficial, for example the absence of NPK fertilizer while maintaining grazing and silage cutting systems.

Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins

P>1. Management of lowland mesotrophic grasslands in north-west Europe often makes use of inorganic fertilizers, high stocking densities and silage-based forage systems to maximize productivity. The impact of these practices has resulted in a simplification of the plant community combined with wide-scale declines in the species richness of grassland invertebrates. We aim to identify how field margin management can be used to promote invertebrate diversity across a suite of functionally diverse taxa (beetles, planthoppers, true bugs, butterflies, bumblebees and spiders). 2. Using an information theoretic approach we identify the impacts of management (cattle grazing, cutting and inorganic fertilizer) and plant community composition (forb species richness, grass species richness and sward architecture) on invertebrate species richness and body size. As many of these management practices are common to grassland systems throughout the world, understanding invertebrate responses to them is important for the maintenance of biodiversity. 3. Sward architecture was identified as the primary factor promoting increased species richness of both predatory and phytophagous trophic levels, as well as being positively correlated with mean body size. In all cases phytophagous invertebrate species richness was positively correlated with measures of plant species richness. 4. The direct effects of management practices appear to be comparatively weak, suggesting that their impacts are indirect and mediated though the continuous measures of plant community structure, such as sward architecture or plant species richness. 5. Synthesis and applications. By partitioning field margins from the remainder of the field, economically viable intensive grassland management can be combined with extensive management aimed at promoting native biodiversity. The absence of inorganic fertilizer, combined with a reduction in the intensity of both cutting and grazing regimes, promotes floral species richness and sward architectural complexity. By increasing sward architecture the total biomass of invertebrates also increased (by c. 60% across the range of sward architectural measures seen in this study), increasing food available for higher trophic levels, such as birds and mammals.

The use of image analysis to estimate population growth rate in Daphnia magna

1. Population growth rate (PGR) is central to the theory of population ecology and is crucial for projecting population trends in conservation biology, pest management and wildlife harvesting. Furthermore, PGR is increasingly used to assess the effects of stressors. Image analysis that can automatically count and measure photographed individuals offers a potential methodology for estimating PGR. 2. This study evaluated two ways in which the PGR of Daphnia magna, exposed to different stressors, can be estimated using an image analysis system. The first method estimated PGR as the ratio of counts of individuals obtained at two different times, while the second method estimated PGR as the ratio of population sizes at two different times, where size is measured by the sum of the individuals' surface areas, i.e. total population surface area. This method is attractive if surface area is correlated with reproductive value (RV), as it is for D. magna, because of the theoretical result that PGR is the rate at which the population RV increases. 3. The image analysis system proved reliable and reproducible in counting populations of up to 440 individuals in 5 L of water. Image counts correlated well with manual counts but with a systematic underestimate of about 30%. This does not affect accuracy when estimating PGR as the ratio of two counts. Area estimates of PGR correlated well with count estimates, but were systematically higher, possibly reflecting their greater accuracy in the study situation. 4. Analysis of relevant scenarios suggested the correlation between RV and body size will generally be good for organisms in which fecundity correlates with body size. In these circumstances, area estimation of PGR is theoretically better than count estimation. 5. Synthesis and applications. There are both theoretical and practical advantages to area estimation of population growth rate when individuals' reproductive values are consistently well correlated with their surface areas. Because stressors may affect both the number and quality of individuals, area estimation of population growth rate should improve the accuracy of predicting stress impacts at the population level.

Towards a population ecology of stressed environments: the effects of zinc on the springtail Folsomia candida

1. To understand population dynamics in stressed environments it is necessary to join together two classical lines of research. Population responses to environmental stress have been studied at low density in life table response experiments. These show how the population's growth rate (pgr) at low density varies in relation to levels of stress. Population responses to density, on the other hand, are based on examination of the relationship between pgr and population density. 2. The joint effects of stress and density on pgr can be pictured as a contour map in which pgr varies with stress and density in the same way that the height of land above sea level varies with latitude and longitude. Here a microcosm experiment is reported that compared the joint effects of zinc and population density on the pgr of the springtail Folsomia candida (Collembola). 3. Our experiments allowed the plotting of a complete map of the effects of density and a stressor on pgr. Particularly important was the position of the pgr= 0 contour, which suggested that carrying capacity varied little with zinc concentration until toxic levels were reached. 4. This prediction accords well with observations of population abundance in the field. The method also allowed us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence. 5. The mechanisms responsible for these phenomena are discussed. As zinc is an essential trace element the initial increase in pgr is probably a consequence of dietary zinc deficiency. The Allee effect may be attributed to productivity of the environment increasing with density at low density. Density dependence is a result of food limitation. 6. Synthesis and applications. We illustrate a novel solution based on mapping a population's growth rate in relation to stress and population density. Our method allows us to demonstrate, simultaneously, hormesis, toxicity, an Allee effect and density dependence in an important ecological indicator species. We hope that the approach followed here will prove to have general applicability enabling predictions of field abundance to be made from estimates of the joint effects of the stressors and density on population growth rate.

The role of satellite image-processing for national-scale estimates of gene flow from genetically modified crops: rapeseed in the UK as a model

1.There is concern over the possibility of unwanted environmental change following transgene movement from genetically modified (GM) rapeseed Brassica napus to its wild and weedy relatives. 2. The aim of this research was to develop a remote sensing-assisted methodology to help quantify gene flow from crops to their wild relatives over wide areas. Emphasis was placed on locating sites of sympatry, where the frequency of gene flow is likely to be highest, and on measuring the size of rapeseed fields to allow spatially explicit modelling of wind-mediated pollen-dispersal patterns. 3. Remote sensing was used as a tool to locate rapeseed fields, and a variety of image-processing techniques was adopted to facilitate the compilation of a spatially explicit profile of sympatry between the crop and Brassica rapa. 4. Classified satellite images containing rapeseed fields were first used to infer the spatial relationship between donor rapeseed fields and recipient riverside B. rapa populations. Such images also have utility for improving the efficiency of ground surveys by identifying probable sites of sympatry. The same data were then also used for the calculation of mean field size. 5. This paper forms a companion paper to Wilkinson et al. (2003), in which these elements were combined to produce a spatially explicit profile of hybrid formation over the UK. The current paper demonstrates the value of remote sensing and image processing for large-scale studies of gene flow, and describes a generic method that could be applied to a variety of crops in many countries. 6.Synthesis and applications. The decision to approve or prevent the release of a GM cultivar is made at a national rather than regional level. It is highly desirable that data relating to the decision-making process are collected at the same scale, rather than relying on extrapolation from smaller experiments designed at the plot, field or even regional scale. It would be extremely difficult and labour intensive to attempt to carry out such large-scale investigations without the use of remote-sensing technology. This study used rapeseed in the UK as a model to demonstrate the value of remote sensing in assembling empirical information at a national level.

Modelling habitat selection and distribution of the critically endangered Jerdon's courser Rhinoptilus bitorquatus in scrub jungle: an application of a new tracking method

1. Jerdon's courser Rhinoptilus bitorquatus is a nocturnally active cursorial bird that is only known to occur in a small area of scrub jungle in Andhra Pradesh, India, and is listed as critically endangered by the IUCN. Information on its habitat requirements is needed urgently to underpin conservation measures. We quantified the habitat features that correlated with the use of different areas of scrub jungle by Jerdon's coursers, and developed a model to map potentially suitable habitat over large areas from satellite imagery and facilitate the design of surveys of Jerdon's courser distribution. 2. We used 11 arrays of 5-m long tracking strips consisting of smoothed fine soil to detect the footprints of Jerdon's coursers, and measured tracking rates (tracking events per strip night). We counted the number of bushes and trees, and described other attributes of vegetation and substrate in a 10-m square plot centred on each strip. We obtained reflectance data from Landsat 7 satellite imagery for the pixel within which each strip lay. 3. We used logistic regression models to describe the relationship between tracking rate by Jerdon's coursers and characteristics of the habitat around the strips, using ground-based survey data and satellite imagery. 4. Jerdon's coursers were most likely to occur where the density of large (>2 m tall) bushes was in the range 300-700 ha(-1) and where the density of smaller bushes was less than 1000 ha(-1). This habitat was detectable using satellite imagery. 5. Synthesis and applications. The occurrence of Jerdon's courser is strongly correlated with the density of bushes and trees, and is in turn affected by grazing with domestic livestock, woodcutting and mechanical clearance of bushes to create pasture, orchards and farmland. It is likely that there is an optimal level of grazing and woodcutting that would maintain or create suitable conditions for the species. Knowledge of the species' distribution is incomplete and there is considerable pressure from human use of apparently suitable habitats. Hence, distribution mapping is a high conservation priority. A two-step procedure is proposed, involving the use of ground surveys of bush density to calibrate satellite image-based mapping of potential habitat. These maps could then be used to select priority areas for Jerdon's courser surveys. The use of tracking strips to study habitat selection and distribution has potential in studies of other scarce and secretive species.

Managing threatened species: the ecological toolbox, evolutionary theory and declining-population paradigm

1. The management of threatened species is an important practical way in which conservationists can intervene in the extinction process and reduce the loss of biodiversity. Understanding the causes of population declines (past, present and future) is pivotal to designing effective practical management. This is the declining-population paradigm identified by Caughley. 2. There are three broad classes of ecological tool used by conservationists to guide management decisions for threatened species: statistical models of habitat use, demographic models and behaviour-based models. Each of these is described here, illustrated with a case study and evaluated critically in terms of its practical application. 3. These tools are fundamentally different. Statistical models of habitat use and demographic models both use descriptions of patterns in abundance and demography, in relation to a range of factors, to inform management decisions. In contrast, behaviour-based models describe the evolutionary processes underlying these patterns, and derive such patterns from the strategies employed by individuals when competing for resources under a specific set of environmental conditions. 4. Statistical models of habitat use and demographic models have been used successfully to make management recommendations for declining populations. To do this, assumptions are made about population growth or vital rates that will apply when environmental conditions are restored, based on either past data collected under favourable environmental conditions or estimates of these parameters when the agent of decline is removed. As a result, they can only be used to make reliable quantitative predictions about future environments when a comparable environment has been experienced by the population of interest in the past. 5. Many future changes in the environment driven by management will not have been experienced by a population in the past. Under these circumstances, vital rates and their relationship with population density will change in the future in a way that is not predictable from past patterns. Reliable quantitative predictions about population-level responses then need to be based on an explicit consideration of the evolutionary processes operating at the individual level. 6. Synthesis and applications. It is argued that evolutionary theory underpins Caughley's declining-population paradigm, and that it needs to become much more widely used within mainstream conservation biology. This will help conservationists examine critically the reliability of the tools they have traditionally used to aid management decision-making. It will also give them access to alternative tools, particularly when predictions are required for changes in the environment that have not been experienced by a population in the past.

Conservation strategy maps: a tool to facilitate biodiversity action planning illustrated using the heath fritillary butterfly

1. The UK Biodiversity Action Plan (UKBAP) identifies invertebrate species in danger of national extinction. For many of these species, targets for recovery specify the number of populations that should exist by a specific future date but offer no procedure to plan strategically to achieve the target for any species. 2. Here we describe techniques based upon geographic information systems (GIS) that produce conservation strategy maps (CSM) to assist with achieving recovery targets based on all available and relevant information. 3. The heath fritillary Mellicta athalia is a UKBAP species used here to illustrate the use of CSM. A phase 1 habitat survey was used to identify habitat polygons across the county of Kent, UK. These were systematically filtered using relevant habitat, botanical and autecological data to identify seven types of polygon, including those with extant colonies or in the vicinity of extant colonies, areas managed for conservation but without colonies, and polygons that had the appropriate habitat structure and may therefore be suitable for reintroduction. 4. Five clusters of polygons of interest were found across the study area. The CSM of two of them are illustrated here: the Blean Wood complex, which contains the existing colonies of heath fritillary in Kent, and the Orlestone Forest complex, which offers opportunities for reintroduction. 5. Synthesis and applications. Although the CSM concept is illustrated here for the UK, we suggest that CSM could be part of species conservation programmes throughout the world. CSM are dynamic and should be stored in electronic format, preferably on the world-wide web, so that they can be easily viewed and updated. CSM can be used to illustrate opportunities and to develop strategies with scientists and non-scientists, enabling the engagement of all communities in a conservation programme. CSM for different years can be presented to illustrate the progress of a plan or to provide continuous feedback on how a field scenario develops.

The joint effects of larval density and C-14-cypermethrin on the life history and population growth rate of the midge Chironomus riparius

1. Chemical effects on organisms are typically assessed using individual-level endpoints or sometimes population growth rate (PGR), but such measurements are generally made at low population densities. In contrast most natural populations are subject to density dependence and fluctuate around the environmental carrying capacity as a result of individual competition for resources. As ecotoxicology aims to make reliable population projections of chemical impacts in the field, an understanding of how high-density or resource-limited populations respond to environmental chemicals is essential. 2. Our objective was to determine the joint effects of population density and chemical stress on the life history and PGR of an important ecotoxicological indicator species, Chironomus riparius, under controlled laboratory conditions. Populations were fed the same ration but initiated at different densities and exposed to a solvent control and three concentrations of C-14-cypermethrin in a sediment-water test system for 67 days at 20 +/- 1 degreesC. 3. Density had a negative effect on all the measured life-history traits, and PGR declined with increasing density in the controls. Exposure to C-14-cypermethrin had a direct negative effect on juvenile survival, presumably within the first 24 h because the chemical rapidly dissipated from the water column. Reductions in the initial larval densities resulted in an increase in the available resources for the survivors. Subsequently, exposed populations emerged sooner and started producing offspring earlier than the controls. C-14-cypermethrin had no effect on estimated fecundity and adult body weight but interacted with density to reduce the time to first emergence and first reproduction. As a result, PGR increased with cypermethrin concentration when populations were initiated at high densities. 4. Synthesis and applications. The results showed that the effects of C-14-cypermethrin were buffered at high density, so that the joint effects of density and chemical stress on PGR were less than additive. Low levels of chemical stressors may increase carrying capacity by reducing juvenile competition for resources. More and perhaps fitter adults may be produced, similar to the effects of predators and culling; however, toxicant exposure may result in survivors that are less tolerant to changing conditions. If less than additive effects are typical in the field, standard regulatory tests carried out at low density may overestimate the effects of environmental chemicals. Further studies over a wide range of chemical stressors and organisms with contrasting life histories are needed to make general recommendations.

Dung and nest surveys: estimating decay rates

1. Wildlife managers often require estimates of abundance. Direct methods of estimation are often impractical, especially in closed-forest environments, so indirect methods such as dung or nest surveys are increasingly popular. 2. Dung and nest surveys typically have three elements: surveys to estimate abundance of the dung or nests; experiments to estimate the production (defecation or nest construction) rate; and experiments to estimate the decay or disappearance rate. The last of these is usually the most problematic, and was the subject of this study. 3. The design of experiments to allow robust estimation of mean time to decay was addressed. In most studies to date, dung or nests have been monitored until they disappear. Instead, we advocate that fresh dung or nests are located, with a single follow-up visit to establish whether the dung or nest is still present or has decayed. 4. Logistic regression was used to estimate probability of decay as a function of time, and possibly of other covariates. Mean time to decay was estimated from this function. 5. Synthesis and applications. Effective management of mammal populations usually requires reliable abundance estimates. The difficulty in estimating abundance of mammals in forest environments has increasingly led to the use of indirect survey methods, in which abundance of sign, usually dung (e.g. deer, antelope and elephants) or nests (e.g. apes), is estimated. Given estimated rates of sign production and decay, sign abundance estimates can be converted to estimates of animal abundance. Decay rates typically vary according to season, weather, habitat, diet and many other factors, making reliable estimation of mean time to decay of signs present at the time of the survey problematic. We emphasize the need for retrospective rather than prospective rates, propose a strategy for survey design, and provide analysis methods for estimating retrospective rates.

Demographic models and the management of endangered species: a case study of the critically endangered Seychelles magpie robin

1. Demographic models are assuming an important role in management decisions for endangered species. Elasticity analysis and scope for management analysis are two such applications. Elasticity analysis determines the vital rates that have the greatest impact on population growth. Scope for management analysis examines the effects that feasible management might have on vital rates and population growth. Both methods target management in an attempt to maximize population growth. 2. The Seychelles magpie robin Copsychus sechellarum is a critically endangered island endemic, the population of which underwent significant growth in the early 1990s following the implementation of a recovery programme. We examined how the formal use of elasticity and scope for management analyses might have shaped management in the recovery programme, and assessed their effectiveness by comparison with the actual population growth achieved. 3. The magpie robin population doubled from about 25 birds in 1990 to more than 50 by 1995. A simple two-stage demographic model showed that this growth was driven primarily by a significant increase in the annual survival probability of first-year birds and an increase in the birth rate. Neither the annual survival probability of adults nor the probability of a female breeding at age 1 changed significantly over time. 4. Elasticity analysis showed that the annual survival probability of adults had the greatest impact on population growth. There was some scope to use management to increase survival, but because survival rates were already high (> 0.9) this had a negligible effect on population growth. Scope for management analysis showed that significant population growth could have been achieved by targeting management measures at the birth rate and survival probability of first-year birds, although predicted growth rates were lower than those achieved by the recovery programme when all management measures were in place (i.e. 1992-95). 5. Synthesis and applications. We argue that scope for management analysis can provide a useful basis for management but will inevitably be limited to some extent by a lack of data, as our study shows. This means that identifying perceived ecological problems and designing management to alleviate them must be an important component of endangered species management. The corollary of this is that it will not be possible or wise to consider only management options for which there is a demonstrable ecological benefit. Given these constraints, we see little role for elasticity analysis because, when data are available, a scope for management analysis will always be of greater practical value and, when data are lacking, precautionary management demands that as many perceived ecological problems as possible are tackled.

Resource availability and the persistence of seed-eating bird populations in agricultural landscapes – a mechanistic modelling approach

1. Reductions in resource availability, associated with land-use change and agricultural intensification in the UK and Europe, have been linked with the widespread decline of many farmland bird species over recent decades. However, the underlying ecological processes which link resource availability and population trends are poorly understood. 2. We construct a spatial depletion model to investigate the relationship between the population persistence of granivorous birds within the agricultural landscape and the temporal dynamics of stubble field availability, an important source of winter food for many of those species. 3. The model is capable of accurately predicting the distribution of a given number of finches and buntings amongst patches of different stubble types in an agricultural landscape over the course of a winter and assessing the relative value of different landscapes in terms of resource availability. 4. Sensitivity analyses showed that the model is relatively robust to estimates of energetic requirements, search efficiency and handling time but that daily seed survival estimates have a strong influence on model fit. Understanding resource dynamics in agricultural landscapes is highlighted as a key area for further research. 5. There was a positive relationship between the predicted number of bird days supported by a landscape over-winter and the breeding population trend for yellowhammer Emberiza citrinella, a species for which survival has been identified as the primary driver of population dynamics, but not for linnet Carduelis cannabina, a species for which productivity has been identified as the primary driver of population dynamics. 6. Synthesis and applications. We believe this model can be used to guide the effective delivery of over-winter food resources under agri-environment schemes and to assess the impacts on granivorous birds of changing resource availability associated with novel changes in land use. This could be very important in the future as farming adapts to an increasingly dynamic trading environment, in which demands for increased agricultural production must be reconciled with objectives for environmental protection, including biodiversity conservation.

The role of management and landscape context in the restoration of grassland phytophagous beetles

1. Declines in area and quality of species-rich mesotrophic and calcareous grasslands have occurred all across Europe.While the European Union has promoted schemes to restore these grasslands, the emphasis for management has remained largely focused on plants. Here we focus on restoration of the phytophagous beetles of these grasslands. Although local management, particularly that which promotes the establishment of host plants, is key to restoration success, dispersal limitation is also likely to be an important limiting factor during the restoration of phytophagous beetle assemblages. 2. Using a 3-year multi-site experiment, we investigated how restoration success of phytophagous beetles was affected by hay-spreading management (intended to introduce target plant species), success in restoration of the plant communities and the landscape context within which restoration was attempted. 3. Restoration success of the plants was greatest where green hay spreading had been used to introduce seeds into restoration sites. Beetle restoration success increased over time, although hayspreading had no direct effect. However, restoration success of the beetles was positively correlated with restoration success of the plants. 4. Overall restoration success of the phytophagous beetles was positively correlated with the proportion of species-rich grassland in the landscape, as was the restoration success of the polyphagous beetles. Restoration success for beetles capable of flight and those showing oligophagous host plant specialism were also positively correlated with connectivity to species-rich grasslands. There was no indication that beetles not capable of flight showed greater dependence on landscape scale factors than flying species. 5. Synthesis and applications. Increasing the similarity of the plant community at restoration sites to target species-rich grasslands will promote restoration success for the phytophagous beetles. However, landscape context is also important, with restoration being approximately twice as successful in those landscapes containing high as opposed to low proportions of species-rich grassland. By targeting grassland restoration within landscapes containing high proportions of species-rich grassland, dispersal limitation problems associated with restoration for invertebrate assemblages are more likely to be overcome.