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.     

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 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.     

Towards a more transparent use of the potential natural vegetation concept – an answer to Chiarucci et al.

In this paper, the concerns of Chiarucci et al. ( 2010 ) regarding use of the potential natural vegetation (PNV) concept are addressed, as voiced in the forum section of the Journal of Vegetation Science. First, we rectify some unfounded expectations concerning PNV, including a relationship with prehuman vegetation and phytosociology. Second, we point out issues that pose considerable challenges in PNV and require common agreement. Here, we address the issue of time and disturbance. We propose to use the static PNV concept as a baseline, a null model for landscape assessment and in comparisons. Instead of changing the PNV concept itself, we introduce a new term, potential future natural vegetation (PFV) to cover estimations of potential successional outcomes. Finally, we offer a new view of PNV with which we intend to make the use of PNV estimates more transparent. We formalize the PNV theory into a partial cause‐effect model of vegetation that clearly states which effects on vegetation are factored out during its estimation. Further, we also propose to assess PNV in a probabilistic setting, rather than providing a single estimate for one location. This multiple PNV would reflect our uncertainty about the vegetation entity that could persist at the locality concerned. Such uncertainty arises from the overlap of environmental preferences of different mature vegetation types. Thus reformulated, we argue that the PNV concept has much to offer as a null model, especially in landscape ecology and in site comparisons in space and time.     

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.     

Ecological palaeoecology in the neotropical Gran Sabana region: Long-term records of vegetation dynamics as a basis for ecological hypothesis testing

Long-term palaeoecological records are needed to test ecological hypotheses involving time, as short-term observations are of insufficient duration to capture natural variability. In this paper, we review the published palaeoecological evidence for the neotropical Gran Sabana (GS) region, to record the vegetation dynamics and evaluate the potential effects of natural climatic and anthropogenic (notably fire) drivers of change. The time period considered (last 13,000 years) covers major global climate changes and the arrival of humans in the region. The specific points addressed are climate–vegetation equilibrium, reversibility of vegetation changes, the origin of extant biodiversity and endemism patterns and biodiversity conservation in the face of global warming. Vegetation dynamics is reconstructed by pollen analysis and fire incidence is deduced from microscopic charcoal records. Palaeoclimatic inferences are derived from global and regional records using independent physico-chemical evidence to avoid circular reasoning. After analyzing all the long-term records available from both GS uplands and highlands, we conclude that: (1) Upland vegetation (mostly treeless savannas and savanna–forest mosaics, with occasional Mauritia palm swamps) is not in equilibrium with the dominant climates, but largely conditioned by burning practices; (2) a hypothetical natural or “original” vegetation type for these uplands has not been possible to identify due to continuous changes in both climate and human activities during the last 13,000 years; (3) at the time scale studied (millennial), the shift from forest to savanna is abrupt and irreversible due to the existence of tipping points, no matter the cause (natural or anthropogenic); (4) on the contrary, the shift from savanna to palm swamps is reversible at centennial time scales; (5) some of the reconstructed past vegetation types have no modern analogues owing to the individual species response to environmental shifts, leading to variations in community composition; (6) extant biodiversity and endemism patterns are not the result of a long history of topographical isolation, as previously proposed but, rather, the consequence of the action of climatic and palaeogeographic variations; (7) the projected global warming will likely exacerbate the expansion of upland savannas by favouring positive fire-climate feedbacks; (8) in the highlands, extinction by habitat loss will likely affect biodiversity but to a less extent that prognosticated by models based only on present-day climatic features; (9) future highland communities will likely be different to present ones due to the prevalence of individual species responses to global warming; and (10) conservation strategies at individual species level, rather than at community level, are enriched by long-term palaeoecological studies analyzed here. None of these conclusions would have been possible to derive from short-term neoecological observations.     

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.     

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.     

The concepts of potential natural vegetation (PNV) and other abstractions (trying to pick up fish with wet hands)

This note follows from an earlier Commentary published in Journal of Biogeography ( Carrión & Fernández, 2009 , 36 , 2202–2203), which provided palaeoecological data, and two replies in the form of Correspondence ( Loidi et al., 2010 , 37 , 2209–2211; Farris et al., 2010 , 37 , 2211–2213). The latter papers attempt to invalidate the palaeoecological database as a source of comparison with the maps of potential vegetation. Here, some of the different interpretations of the term ‘potential natural vegetation’ (PNV), as used by the floristic phytosociological school, are discussed. It is suggested that there is a conceptual impasse that will not have a solution until a terminological consensus is reached. This terminology will open new methodological avenues that will facilitate the entry of new information derived from historical biogeography, palaeoecology, ecology, phylogeography, and niche and community modelling. One of the main sources of conflict arises from the link made between habitats and floristically‐determined associations, a confusion that has crucial repercussions in biological conservation, including in respect of the EU Habitats Directive.     

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.     

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.     

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.     

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.     

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.     

Reconstructing pre-impact vegetation cover in modified landscapes using environmental modelling, historical surveys and remnant vegetation data: a case study in the Fleurieu Peninsula, South Australia

Aim To investigate the application of environmental modelling to reconstructive mapping of pre‐impact vegetation using historical survey records and remnant vegetation data. Location The higher elevation regions of the Fleurieu Peninsula region in South Australia were selected as a case study. The Fleurieu Peninsula is an area typical of many agricultural regions in temperate Australia that have undergone massive environmental transformation since European settlement. Around 9% of the present land cover is remnant vegetation and historical survey records from the ad 1880s exist. It is a region with strong gradients in climate and topography. Methods Records of pre‐impact vegetation distribution made in surveyors’ field notebooks were transcribed into a geographical information system and the spatial and classificatory accuracy of these records was assessed. Maps of remnant vegetation distribution were obtained. Analysis was undertaken to quantify the environmental domains of historical survey record and remnant vegetation data to selected meso‐scaled climatic parameters and topo‐scaled terrain‐related indices at a 20 m resolution. An exploratory analytical procedure was used to quantify the probability of occurrence of vegetation types in environmental domains. Probability models spatially extended to geographical space produce maps of the probability of occurrence of vegetation types. Individual probability maps were combined to produce a pre‐impact vegetation map of the region. Results Surveyors’ field notebook records provide reliable information that is accurately locatable to levels of resolution such that the vegetation data can be spatially correlated with environmental variables generated on 20 m resolution environmental data sets. Historical survey records of vegetation were weakly correlated with the topo‐scaled environmental variables but were correlated with meso‐scaled climate. Remnant vegetation records similarly not only correlated to climate but also displayed stronger relationships with the topo‐scaled environmental variables, particularly slope. Main conclusions A major conclusion of this study is that multiple sources of evidence are required to reconstruct past vegetation patterns in heavily transformed region. Neither the remnant vegetation data nor historical survey records provided adequate data sets on their own to reconstruct the pre‐impact vegetation of the Fleurieu Peninsula. Multiple sources of evidence provide the only means of assessing the environmental and historical representativeness of data sets. The spatial distribution of historical survey records was more environmentally representative than remnant vegetation data, which reflect biases due to land clearance. Historical survey records were also shown to be classificatory and spatially accurate, thus are suitable for quantitative spatial analyses. Analysis of different spatial vegetation data sets in an environmental modelling framework provided a rigorous means of assessing and comparing respective data sets as well as mapping their predicted distributions based on quantitative correlations. The method could be usefully applied to other regions where predictions of pre‐impact vegetation cover are required.     

Role of disturbed vegetation in mapping the boreal zone in northern Eurasia

Question: Is there a need for disturbance mapping integrated in the CircumBoreal Vegetation Mapping Program? Location: Eurasian boreal forest. Disturbance and mapping: The boreal zone is characterized by a multitude of natural and anthropogenic disturbance agents with importance over a wide range of spatial and temporal scales. Disturbance is a prime driver of succession in most of the boreal zone, producing landscape diversity characterized by a large‐scale vegetation mosaic of early to late succession states. When mapping the circumboreal vegetation, spatial extent, time involved from disturbance to recovered condition and likelihood of interacting disturbance types are crucial for how current vegetation is interpreted and subsequently included as map characteristics. In this paper we present examples from the boreal zone where natural and/or anthropogenic disturbance regimes dominate the state and distribution of vegetation, and possibilities for assessing the nature and extent of the disturbed regions using remotely sensed data. Conclusion: Disturbed vegetation occupies large areas in the boreal zone and related vegetation successions should be adequately represented when mapping the zone. In regions where the ‘potential natural vegetation’ is a hypothetical reconstruction from remnants of ‘natural’ vegetation it would be preferable to use the concept of ‘actual real vegetation’ for which remote sensing at coarse, medium and fine resolution is an efficient tool. The Land Cover Classification System (LCCS) may offer sufficient flexibility to incorporate information about the disturbance of circumboreal vegetation.     

Bioclimatic classification of US vegetation along a coast-to-coast macrotransect crossing central United States. I: Mediterranean vegetation

This report presents the first of two parts of a bioclimatic classification of the vegetation of the United States. Using a geographical information system, 987 weather stations were located along a longitudinal macrotransect from the shores of the Atlantic to Pacific on four maps: Map of the Physiographic Divisions of the Conterminous US, US Potential Natural Vegetation Map, US Ecoregion Map, and Terrestrial Ecosystems-Isobioclimates Map of the Conterminous United States. Based on these maps, bibliographic resources and field data, we deduced the potential natural vegetation (PNV) of each weather station; then, we assigned the different PNV types to alliance or association levels using the US National Vegetation Classification (USNVC). In a next step, USNVC groups were related with similar level phytosociological syntaxa described in the study area. The bioclimatic distribution of the USNVC units defined was then interpreted using the bioclimatic classification proposed in successive approximations by S. Rivas-Martínez. The distribution of USNVC alliances was mainly linked to the macrobioclimates (Mediterranean, Temperate, and Tropical) of the longitudinal gradient examined, though some edaphic factors induced the appearance of specialized plant groups. Herein, we present our data for the Mediterranean macrobioclimate, in which 53 alliances and 28 isobioclimates were identified.     

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.     

Retrospective monitoring of rangeland vegetation change: ecohistory from deposits of sheep dung associated with shearing sheds

This paper explores the potential of a new method of reconstructing historical vegetation change in the Australian rangelands. Historical monitoring of rangeland vegetation has been so deficient that it is not possible to determine whether a long‐term trend toward degradation has occurred (as is often assumed) or, indeed, if it is continuing to occur. Because long‐term records are unavailable any attempt to monitor vegetation retrospectively must be based on proxy measures rather than direct observation. Where historical data are lacking an integration of palaeoecological, archaeological and ecological methods is required to reconstruct the past. Our research is based on a detailed analysis of sheep faeces deposited near a shearing shed in the semiarid rangelands of south‐west Queensland between the late 1930s and the mid‐1990s. The faeces in these deposits represent the diet of sheep in the days leading up to the property’s annual shearing and as such are a potentially useful index to changes in vegetation. Results indicate significant changes in the diet of sheep since the late 1940s. The potential of this method, and its limitations, are discussed. Long‐term records are critical in understanding issues of sustainability in land management and it is intended that this paper will stimulate further research into historical vegetation change in rangelands.