2003 Kluwer Academic Publishers. Printed in the Netherlands.
223 The implication of past and present landscape patterns for biodiversity research: introduction and overview Isabelle Poudevigne 1 & Jacques Baudry 2 1 Landscape Systems Research Group, Facult e des Sciences, Universit e de Rouen, 76821 Mont Saint Aignan, (E- mail: Isabelle.Poudevigne@univ-rouen.fr) 2 INRA SAD Armorique, CS 84215, 65 Rue de Saint Brieuc, 35042 Rennes C edex, France Since its development in the early 1980s, the aim of landscape ecology is to understand both the effects of spatial patterns on ecological processes and the development of those spatial patterns. In Europe, hu- man activities have had a pronounced long term role in shaping this heterogeneity so that Europe is now the most modied of any continent (Godron and For- man 1983). Therefore, research on how humans have shaped landscapes is of utmost importance (Burel and Baudry 1999). If changes following the World War II are a frequent topic, long term changes are also impor- tant (Berlung 1991) as they have, in interaction with the physical environment, selected the regional species pool. Indeed, present co-existence of species in the landscape is thought to be the end product of a series of long term responses between species requirements and environmental gradients of both natural and hu- man origin (Huston 1994). Despite potential adverse effects, many long-term human disturbance regimes have promoted biodiversity. In western Europe, tra- ditional low intensity agriculture has often promoted high levels of diversity, or provided relict habitat for rare or threatened taxa (Green 1990). By analysing ecological patterns and functioning at large spatial and temporal scale, landscape ecology offers some of the most successful examples of the uptake of ecology in policy or management (Ormerod et al. 2002). If re- gional approaches are increasing (Surez-Seoane et al. 2002), landscape studies of a lesser extent (few km 2 ) are still dominant. One of the most important current areas in which the study of landscape patterns, past and present, offers a major path of research is in understanding the sustain- able management of biodiversity. This was the theme of a meeting of landscape ecologists held in Rouen, France in October 2001. Our aim with this array of selected papers from the meeting is to share our out- look on current European trends and approaches to landscape ecology. Patterns of human activities in landscapes, what heritage? Cubizolle et al. (2003) give evidence that human ac- tivities, as early as the Iron age in France, had an important inuence on plant biodiversity whose reec- tion we see today. Their analysis of mires in France, based on radiocarbon dates and pollen analysis, reveal that some activities such as the building of small dams, forest clearing, and cattle grazing have favoured peat inception. Ernoult et al. (2003) provide an analysis of changes in agricultural landscapes during the last decades. They do so by proposing two measures of landscape organisation, i.e., how patterns deviate from randomness. The rst measure deals with the spatial relationships among the different categories of land use intensity that form the landscape mosaic. The sec- ond measure assess the relationships between land use patches and the physical environment. They demon- strate that changes in agriculture have led to more stochastic patterns. Assessing the risks to biodiversity This historical background explains why today many representative or threatened species are dispersed through environments, such as farmland, that are in economic use (Robinson and Sutherland 2002). Un- fortunately, recent changes in agricultural manage- ment and intensication have altered subtle equilibria between biota and patterns of use in both farmed and natural landscapes even on areas with strong constraints to development (Poudevigne et al. 2002). 224 Habitat alteration (including habitat loss, degradation and fragmentation) is now among the major risks of ecosystem degradation by these human activities (Whiteld et al. 2002). Many of these empirical stud- ies seek the variable that at different scales, from patch to landscapes drive species distribution. Land- scape ecologists have proposed many measures to assess the relationship between spatial heterogeneity and biodiversity in landscapes (Cullinan and Thomas 1992; McGarigal and Marks 1995), but few effec- tively relate pattern to process (Levin 1992). Classical metrics remain dissatisfying as correlation between species diversity indices and heterogeneity indices are not always meaningful. A step further is to identify the thresholds to which ecosystems can be modied without being irreversibly altered (Suter 1993). Jeanneret et al. (2003) compare the distribution of carabids, spiders and butteries in two Swiss agri- cultural landscapes. They conclude that landscape metrics are of little use, while information on habitat mosaic provide information. It also appears that the variables explaining most of the variance in species distribution are different in the two landscapes, as are the reactions of the different groups of species. This work calls for comparative studies along land- scape gradients. Milln de la Pea et al. (2003) found little correlation between the landscape descriptors they investigated, such as the amount of cropland, and the richness and species composition of small mammal communities on agricultural land in western France. They nevertheless found effects on mammal demography: intensication of agriculture had re- duced the density of rare and habitat specialist species while favouring habitat-generalists. Mennechez et al. (2003), investigating on the effects of habitat loss and fragmentation on buttery population functioning with classical measures, argue that spatial heterogeneity in their case study affects dispersal more than demog- raphy. With this organism-centred point of view, the authors then propose the denition of a new parame- ter, the minimal patch area needed to establish a local population in highly fragmented landscapes. Looking beyond France, in Africa, Fritz et al. (2003) observe that the extension of agriculture along rivers in the Mid Zambezi valley, Zimbabwe, impacts on most wild species. But they also dene a thresh- old value of eld size above which there seems to be an acceleration of the decrease in wildlife density and diversity. Denition of such thresholds may be an important asset in determining priorities for the management of degraded ecosystems. Modelling and eld testing the risks Rather than synthesising the landscape system with metrics, some scientists have chosen to model these systems, in an attempt to capture manageable aspects of their complexity that are often beyond eld exper- imentation and assessment (Jaberg and Guisan 2002). Baudry et al. (2003) model dairy farm landscapes with a measure which both considers the landscape and the species. Connectivity is described as a measure of landscape structure and species characteristics based on individual area requirements and dispersal distance. Results reveal that for one farming system, landscape connectivity remains the same over years (in a 7-year model experiment), while it is signicantly differ- ent between two intensive and traditionally extensive farming systems). This work also suggests the im- portance of considering the temporal dimensions of spatial heterogeneity metrics. Similarly, Cousins et al. (2003) present a model which explores the effects of grazing frequency and intensity on plant persistence, and the relative effects of grassland size and pattern. These models aim at exploring the effects of further fragmentation and habitat loss on the persistence of species or plant functional groups. This generation of models aim to be valuable tools for managers, both to dene threshold risk values, and do simulations of the potential impacts of landscape planning decisions. Most landscape ecology research is based on a cor- relative approach, mainly because of the scale and complexity of the landscape systems. Large scale and in vivo experimentation, because it is either unaf- fordable or technically unfeasible, is rarely, with few exceptions (Bradley and Ormerod 2002; Donlan et al. 2002), a possible way of testing hypotheses proposed by the correlative studies that often characterize land- scape ecology. Equally, landscape ecologists have so far rarely moved beyond the realms of pattern among species and communities. Yet, in that eld, analy- sis of spatial genetic structuring can yield interesting possibilities. Arnaud et al. (2003), working the land snail Helix aspersa, test landscape-based geographical distance to an isolation by distance model. 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