Quantitative comparison of the diversity of landscapes with actual vs. potential natural vegetation

Abstract. In the past 20 years, several metrics have been developed to quantify various aspects of landscape structure and diversity in space and time, and most have been tested on grid‐based thematic maps. Once landscape patterns have been quantified, their effects on ecological functions can be explained if the expected pattern in the absence of specific processes is known. This type of expected pattern has been termed a neutral landscape model. In the landscape‐ecological literature, researchers traditionally adopt random and fractal computer‐generated neutral landscape models to verify the expected relationship between a given ecological process and landscape spatial heterogeneity. Conversely, little attention has been devoted to distribution patterns of potential natural vegetation (PNV) as an ecological baseline for the evaluation of pattern‐process interactions at the landscape scale. As an application for demonstration, we propose a neutral model based on PNV as a possible reference for a quantitative comparison with actual vegetation (AC V) distribution. Within this context, we introduce an evenness‐like index termed ‘actual‐to‐potential entropy ratio’ (H_A/P = H_ACV/H_PNV, where H is Shannon's entropy). Results show that, despite the hypothetical character of most PNV maps, the use of PNV distribution as a baseline for a quantitative comparison with ACV distribution may represent a first step towards a general model for the evaluation of the effects of disturbance on vegetation patterns and diversity.     

Potential natural vegetation: validity and applicability in landscape planning and nature conservation

Abstract. Since the introduction of ‘potential natural vegetation’ (PNV) as a concept in vegetation science by Tüxen (1956), many PNV-maps with different scales have been made. Tüxen emphasized the great value of PNV-maps for different purposes in land use, landscape planning and nature conservation, in particular with regard to forestry, agriculture and landscape management. Different aspects are discussed in order to examine the validity and applicability of PNV-maps in landscape planning and nature conservation. PNV-maps are useful for the differentiation of natural and landscape units on a small scale (< 1 : 100 000). However, maps of the potential natural vegetation are less useful for purposes of detailed planning on larger scales (> 1 : 100 000). Problems arise, for example, from the often highly hypothetical character of the construction and the practice of taking remnants of ‘natural’ vegetation as a reference object for the PNV. With regard to the goals of modern landscape planning and nature conservation purposes (e.g. conserving biodiversity in the cultural landscape of Central Europe) the exact documentation of the actual real vegetation (ARV) on intermediate and large scales gives much more detailed information than a hypothetical PNV.     

The power of potential natural vegetation (and of spatial‐temporal scale): a response to Carrión & Fernández (2009)

A commentary by Carrión & Fernández (Journal of Biogeography, 2009, 36 , 2202–2203) compared Holocene pollen records with models of potential natural vegetation (PNV) proposed in the phytosociological literature and concluded that the predicted PNV resulted from anthropogenic disturbance. However, the authors misinterpreted PNV, leading to two serious flaws in their assumptions: (1) PNV is not defined as a pre‐anthropic or climax plant community; and (2) PNV is not a concept restricted to the phytosociological method. Therefore we criticize the conclusions expressed in the commentary, and we stress the need for an interdisciplinary approach based on multi‐temporal and multi‐spatial scales to achieve a modern framework for the study of plant communities.     

Traditional land uses enhanced plant biodiversity in a Mediterranean agro-silvo-pastoral system

Mediterranean agro-silvo-pastoral systems play a key role in view of the positive contribution that they could offer to a sustainable development of European agriculture. The knowledge of the vegetation dynamics and of the processes and land uses favoring different vegetation types related to the same actual potential natural vegetation (PNV) could represent a sound reference framework for monitoring and managing plant biodiversity in these systems. The aim of the research was to evaluate plant diversity along a gradient of use intensity comparing the actual vegetation versus the PNV. The results of our research showed that in the studied Mediterranean agro-silvo-pastoral system, included in the same environmental unit, human activities enhanced plant biodiversity. Moreover, the case study presented here confirmed the effectiveness of those landscape approaches comparing actual vegetation versus the PNV for plant biodiversity monitoring and reinforced previous studies showing the effect of human activities on plant community diversity at the environmental unit scale in different biogeographical contexts.     

Long‐term vegetation stability and the concept of potential natural vegetation in the Neotropics

When vegetation trends over time are analysed from an appropriate long‐term perspective using palaeoecological records, the concept of potential natural vegetation (PNV) is unsupported because of continual vegetation changes driven by natural or anthropic forcings. However, some palaeoecological records show long‐lasting (i.e. millennial) vegetation stability at multidecadal to centennial time scales in the absence of natural and human drivers of change, which fits within the definition of PNV. A more detailed palaeoecological analysis of these records shows that they are an exception rather than a rule, and that they cannot be differentiated from other transient ecological states. Therefore, long records of vegetation stability cannot be considered to be valid evidence for PNV. From a palaeoecological perspective, the PNV concept is concluded to be unnecessary, even in cases of multidecadal to centennial vegetation stability in the absence of environmental disturbance.     

Understanding properly the `potential natural vegetation' concept

This is a response to critical comments concerning the inappropriate use of the potential natural vegetation (PNV) concept made in a recent contribution to the Commentary section of this journal. We consider that the PNV concept has been misinterpreted. PNV has been used extensively in several European countries since the mid‐1950s and was never intended to be used to make a prediction of what vegetation would dominate in an area if human influence were removed. PNV maps express hypothetical assumptions of what corresponds to dominant or natural vegetation in each area. Remnants of the vegetation of the past provided by palaeopalynology and other disciplines provide valuable information for interpreting modern vegetation, but natural changes and anthropogenic influences operating over the last millennia have to be taken into account. Annex I of the Habitats Directive provides a balanced list of habitat types for implementing conservation policies within the European Union.     

Potential natural vegetation and pre‐anthropic pollen records on the Azores Islands in a Macaronesian context

This paper discusses the concept of potential natural vegetation (PNV) in the light of the pollen records available to date for the Macaronesian biogeographical region, with emphasis on the Azores Islands. The classical debate on the convenience or not of the PNV concept has been recently revived in the Canary Islands, where pollen records of pre‐anthropic vegetation seemed to strongly disagree with the existing PNV reconstructions. Contrastingly, more recent PNV model outputs from the Azores Islands show outstanding parallelisms with pre‐anthropic pollen records, at least in qualitative terms. We suggest the development of more detailed quantitative studies to compare these methodologies as an opportunity for improving the performance of both. PNV modelling may benefit by incorporating empirical data on past vegetation useful for calibration and validation purposes, whereas palynology may improve past reconstructions by minimizing interpretative biases linked to differential pollen production, dispersal and preservation.     

Application of the Ancient Forest Concept to Potential Natural Vegetation Mapping in Flanders, A Strongly Altered Landscape in Northern Belgium

Construction of potential natural vegetation (PNV) poses particular challenges in landscapes heavily altered by human activity and must be based on transparent, repeatable methods. We integrated the concept of ancient forest (AF) and ancient forest species (AFS) into a four-step procedure of PNV mapping: 1) classification of forest vegetation relevés; 2) selection of those vegetation types that can serve as PNV units, based on AF and AFS; 3) merging of selected vegetation types into five PNV units that can be predicted from a digital morphogenetic soil map; 4) mapping of three additional PNV units based on additional environmental data. The second step, concerning the selection of reference forest vegetation, is of particular interest for PNV construction in Flanders (northern Belgium), where forest cover has been subject to temporal disruption and spatial fragmentation. Among the variety of extant forest recovery states, we chose as PNV units those vegetation types for which a high proportion of relevés had been located in AF and that contained many AFS. As the frequency of AFS depends on site conditions, we only compared and selected vegetation types that are found on similar sites according to average Ellenberg indicator values. While succession is irrelevant for the definition of PNV, colonization rates of AFS can be used to estimate the time required for PNV to be restored in a site.     

Improving Transboundary Maps of Potential Natural Vegetation Using Statistical Modeling Based on Environmental Predictors

The potential natural vegetation (PNV) is a tool for landscape planning, nature preservation and the assessment of naturalness. It is mostly constructed by the application of expert knowledge. This paper shows the advantages of using a more sophisticated and formalized PNV construction that overlays vegetation types and site factor maps by applying a Bayes model and herewith improving existing PNV maps solely based on expert knowledge. The investigation was conducted in the forest complex of the Bavarian Forest National Park (Germany) and the adjacent ?umava National Park (Czech Republic). The project reached two major results: (1) The existing heterogeneous country-specific databases of natural site conditions and of vegetation types could be adapted to each other to construct a solid scientific basis to deduce a PNV map. The habitat requirements of the occurring harmonized vegetation types can now be quantitatively described in a formalized way. (2) The combination of terrestrial PNV mapping and numerical modeling allows the synthesis of the views of the different experts that generated the maps used for model calibration. However, the modeled map loses the details of the expert-based map that cannot be derived from the underlying site maps. A common modeled PNV map of both national parks covering an area of about 92,000 ha was created. While the former expert-based PNV maps display breaks along the country border, the modeled PNV presents a harmonized view based on the common database of both national parks.     

On the theoretical concept of the potential natural vegetation and proposals for an up-to-date modification

In the extent to which it is used, the concept of the potential natural vegetation (PNV) is one of the most successful novelties in vegetation science over the last decades. However, previous applications of the concept have shown that the theoretical principles were used inconsistently or interpreted in an incorrect sense. The present problems in application (which become evident when visualizing historical aspects of the concept) mainly result from (a) inconsistent treatment of the construction criteria; (b) failure to distinguish between the “potential natural vegetation”, the “reconstructed natural vegetation” and the vegetation developing during succession, (c) the lack of a precise definition for reference terms to construct potential natural vegetation (e.g. treating reversible vs. irreversible changes of vegetation). For a sensible application of the concept it is suggested (a) to construct the potential natural vegetation on the basis of natural site conditions as well as permanently effective site changes as a consequence of human impact, (b) to consider the PNV to be in balance with all site conditions taken as basis for its construction. In practice, however, the construction basis may also derive from a particular question underlying the making of a PNV-map. A suggestion for a re-definition of the term “potential natural vegetation” as well as a key for PNV-mapping (valid for landscapes of Northern Germany) are given.     

Natural,potential and actual vegetation in North America

The potential natural vegetation (PNV) concept has parallel applications in Europe and North America. Paleoecological studies in parts of North America provide records of vegetation patterns and dynamics under little or no human disturbance. Something resembling PNV emerges at millennial temporal scales and at regional to subcontinental spatial scales. However, at finer spatial and temporal scales, actual vegetation often displays properties of inertia, contingency and hysteresis, most frequently because of climatic variability across multiple timescales and the episodic nature of disturbance and establishment. Thus, in the absence of human disturbance, the actual vegetation that develops at a site may not resemble a particular PNV ideal, but could instead represent one of any number of potential outcomes constrained by historically contingent processes. PNV may best be viewed as an artificial construct, with utility in some settings. Its utility may diminish and even be detrimental in a rapidly changing environment.     

The concept of potential natural vegetation: an epitaph?

We discuss the usefulness of the concept of Potential Natural Vegetation (PNV), which describes the expected state of mature vegetation in the absence of human intervention. We argue that it is impossible to model PNV because of (i) the methodological problems associated to its definition and (ii) the issues related to the ecosystems dynamics.We conclude that the approach to characterizing PNV is unrealistic and provides scenarios with limited predictive power. In places with a long‐term human history, interpretations of PNV need to be very cautious, and explicit acknowledgement made of the limitations inherent in available data.     

Potential replacement vegetation: an approach to vegetation mapping of cultural landscapes

Abstract. The concept of mapping potential replacement vegetation (PRV) is proposed as a parallel to potential natural vegetation (PNV). Potential replacement vegetation (PRV) is an abstract and hypothetical vegetation which is in balance with climatic and soil factors currently affecting a given habitat, with environmental factors influencing the habitat from outside such as air pollution, and with an abstract anthropogenic influence (management) of given type, frequency and intensity. For every habitat, there is a series of possible PRV-types corresponding to the different anthropogenic influences, e.g. grazing, mowing, trampling or growing cereals. The PRV-concept is especially useful in large-scale mapping (scales > 1 : 25 000) of small areas where replacement vegetation is the focus of attention for managers and land-use planners, for example in nature reserves where the aim is conservation of replacement vegetation managed in a traditional way, or in restoration ecology where the concept may be used for defining restoration goals and evaluating the success of restoration efforts. At smaller scales, PRV-mapping may be useful for revealing the biogeographical patterns of larger areas which may be different from the corresponding PNV patterns, because replacement vegetation and natural vegetation may respond to environmental gradients at different scales. An example of medium-scale PRV-mapping through the coincidence of diagnostic species of vegetation types, based on species distribution grid data, is presented. In cultural landscapes, the advantage of using the PRV-concept instead of PNV is its direct relationship to the replacement vegetation. In the habitat mapping with respect to the replacement vegetation, the PRV concept yields more valuable results than the mapping of actual vegetation, as the latter is strongly affected by spatially variable anthropogenic influences which may be largely independent from climatic and soil factors.     

Precise vegetation restoration in arid regions based on potential natural vegetation and potential normalized difference vegetation index

Precise vegetation restoration is critical in drylands, as some inappropriate restoration attempts have even increased water scarcity and degradation in afforestation areas. Potential natural vegetation (PNV) is widely used to provide a reference for the appropriate location and vegetation type of restoration programs while the appropriate restored areas remain unknown. Therefore, we proposed a PNV–potential normalized difference vegetation index (PNDVI) coupling framework based on multiple machine learning (ML) algorithms for precise dryland vegetation restoration. Taking the lower Tarim River Basin (LTRB) with a total area of 1,182 km² as a case study, its present suitable restoration locations, area, and appropriate planting species were quantitatively estimated. The results showed that the model developed by incorporating PNDVI into PNV with easily measurable and available data such as temperature and soil properties can accurately identify dryland restoration patterns. In LTRB, the potentially suitable habitats of trees and grass are closer to the riverbank, while shrubby habitats are further away from the course, covering 1.88, 2.96, and 25.12 km², respectively. There is still enormous land potential for further expansion of the current trees and grass in the LTRB, with 2.56 and 1.54% of existing land supposed to be trees and grass, respectively. This study's novel aspect is combining PNV and PNDVI to quantify and estimate precise restoration patterns through multiple ML algorithms. The model developed here can be used to evaluate the suitable reforestation locations, area, and vegetation types in drylands and to provide a basis for precise vegetation restoration.     

Are potential natural vegetation maps a meaningful alternative to neutral landscape models?

Abstract. In this paper, we present a short overview of neutral landscape models traditionally adopted in the landscape ecological literature to differentiate landscape patterns that are the result of simple random processes from patterns that are generated from more complex ecological processes. Then, we present another family of models based on Tuxen’ s definition of potential natural vegetation that play an important role, especially in Europe, for landscape planning and management. While neutral landscape models by their very nature do not take into account vegetation dynamics, nor abiotic constraints to vegetation distribution, the concept of potential natural vegetation includes the effects of vegetation dynamics in a spatially explicit manner. Therefore, we believe that distribution maps of potential natural vegetation may represent an ecological meaningful alternative to neutral landscape models for evaluating the effects of landscape structure on ecological processes.     

Predictive modeling of the potential natural vegetation pattern in northeast China

Based on the characteristics of natural vegetation distribution in northeast China, using multivariate analysis and geographical information system technology, we established a regional ‘vegetation–environment’ model to simulate geographical distribution of 16 natural vegetation types under present environmental conditions, representing the potential natural vegetation (PNV) of northeast China, on the basis of digital maps of seven environmental variables including climate and topography. Comparison of simulated PNVs distributions with the actual natural vegetation distribution indicated a good agreement, with overall predictive accuracy of 66.9% and overall Kappa value of 0.67. The predictions of model, however, were poor, for only 0.62 of AUC value was yielded. The current resolution and accuracy of the model can be applied to simulate and map the natural vegetation pattern at the regional scale and also used to analyze the effect of climatic changes on natural vegetation.     

Dealing with uncertainty in landscape genetic resistance models: a case of three co‐occurring marsupials

Landscape genetics lacks explicit methods for dealing with the uncertainty in landscape resistance estimation, which is particularly problematic when sample sizes of individuals are small. Unless uncertainty can be quantified, valuable but small data sets may be rendered unusable for conservation purposes. We offer a method to quantify uncertainty in landscape resistance estimates using multimodel inference as an improvement over single model‐based inference. We illustrate the approach empirically using co‐occurring, woodland‐preferring Australian marsupials within a common study area: two arboreal gliders (Petaurus breviceps, and Petaurus norfolcensis) and one ground‐dwelling antechinus (Antechinus flavipes). First, we use maximum‐likelihood and a bootstrap procedure to identify the best‐supported isolation‐by‐resistance model out of 56 models defined by linear and non‐linear resistance functions. We then quantify uncertainty in resistance estimates by examining parameter selection probabilities from the bootstrapped data. The selection probabilities provide estimates of uncertainty in the parameters that drive the relationships between landscape features and resistance. We then validate our method for quantifying uncertainty using simulated genetic and landscape data showing that for most parameter combinations it provides sensible estimates of uncertainty. We conclude that small data sets can be informative in landscape genetic analyses provided uncertainty can be explicitly quantified. Being explicit about uncertainty in landscape genetic models will make results more interpretable and useful for conservation decision‐making, where dealing with uncertainty is critical.     

Model for the potential natural vegetation mapping of Friuli Venezia-Giulia (NE Italy) and its application for a biogeographic classification of the region

ABSTRACT

The aim of this paper is to present a joint vegetation data base and GIS application to produce a model to map the potential natural vegetation (PNV) of the Friuli-Venezia Giulia region (NE Italy) and to show how the map can be used to draw a biogeographic classification of the region. All the natural arboreal coenoses growing below the timber line, as well as the dwarf shrubs and prairies developing above this limit, were considered. Some cross sections, extracted from the potential vegetation map, were tested against transects of real vegetation distribution.     

The TRM Model of Potential Natural Vegetation in Mountain Forests

Due to advances in spatial modeling and improved availability of digital geodata, traditional mapping of potential natural vegetation (PNV) can be replaced by ecological modeling approaches. We developed a new model to map forest types representing the potential natural forest vegetation in the Bavarian Alps. The TRM model is founded on a three-dimensional system of the ecological gradients temperature (T), soil reaction (R), and soil moisture (M). Within such a “site cube” forest types are defined as homogenous site units that give rise to forest communities with comparable species composition, structure, production and protective functions. The three gradients were modeled using regression algorithms with area-wide, high resolution geodata on climate, relief and soil as predictors and average Ellenberg indicator values for temperature, acidity and moisture of vegetation plots as dependent variables summarizing plant responses to ecological gradients. The resulting predictor-response relationships allowed us to predict gradient positions of each raster cell in the region from geodata layers. The three-dimensional system of gradients was partitioned into 26 forest types, which can be mapped for the whole region. TRM-based units are supplemented by 22 forest types of special sites defined by other ecological factors such as geomorphology, for which individual GIS rules were developed. The application of our model results in an intermediate-scale map of potential natural forest vegetation, which is based on an explicit function of temperature, reaction and moisture and is therefore consistent and repeatable in contrast to traditional PNV maps.