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.     

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.     

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.     

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.     

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.     

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.     

Plant Communities in Multi-Metal Contaminated Soils: A Case Study in the National Park of Alta Murgia (Apulia Region - Southern Italy)

A phytosociological study was conducted in the National Park of Alta Murgia in the Apulia region (Southern Italy) to determine the adverse effects of metal contamination of soils on the distribution of plant communities. The phytosociological analyses have shown a remarkable biodiversity of vegetation on non-contaminated soils, while biodiversity appeared strongly reduced on metal-contaminated soils. The area is naturally covered by a wide steppic grassland dominated by Stipa austroitalica Martinovský subsp. austroitalica. Brassicaceae such as Sinapis arvensis L. are the dominating species on moderated contaminated soils, whereas spiny species of Asteraceae such as Silybum marianum (L.) Gaertn. and Carduus pycnocephalus L. subsp. pycnocephalus are the dominating vegetation on heavily metal-contaminated soils. The presence of these spontaneous species on contaminated soils suggest their potential for restoration of degraded lands by phytostabilization strategy.     

Vegetation of the dry bioclimatic areas in the Dominican Republic

We conduct a study of the dry forest in areas of scant rainfall in the Dominican Republic; these are sites with particular endemic habitats, as the samples reveal a high rate of local and insular endemic species, with 84 (24.85%) endemisms. This work covers dry forests rich in coarse spiny plants and includes a statistical (multivariate analysis), phytosociological and catenal study of the vegetation. We analyse the floristic composition, ecology, distribution and richness in endemisms of each association. The bioclimatic analysis reveals the different thermotypes and ombrotypes on the island, and locates the proposed plant associations in the infra- and thermotropical thermotypes, and in the arid, semiarid and dry ombrotypes. As a result of this phytosociological analysis, we propose the new alliance Harrio nashii–Acacion skleroxylae and four plant associations: Harrisio nashii–Prosopidetum juliflorae, Crotono poitaei–Erythroxyletum rotundifolii, Lonchocarpo pycnophylli–Cylindropuntietum caribaeae and Neoabbottio paniculatae–Guaiacetum officinali; these associations connect catenally with the subhumid forest and mangrove swamps.     

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.     

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.     

The vegetation of alpine belt karst-tectonic basins in the central Apennines (Italy)

Abstract

The vegetation communities of the karst-tectonic basins of the Majella massif alpine belt were studied using the phytosociological methods, and analysed from coenological, synchorological and syntaxonomical viewpoints. During the field-work, 115 relevés were performed using the phytosociological approach of Braun-Blanquet, and these relevés were further subjected to multivariate analyses. Eight clusters of relevés resulted from the numerical classification. The plant communities identified in the study area were ascribed to the following five associations, two sub-associations and one community type: Leontopodio – Seslerietum juncifoliae (ass. nova); Helianthemo – Festucetum italicae (ass. nova); Gnaphalio – Plantaginetum atratae; Taraxaco – Trifolietum thalii gnaphalietosum magellensis (subass. nova); Luzulo italicae – Nardetum, Carici – Salicetum retusae; Saxifrago – Papaveretum julici, Saxifrago – Papaveretum androsacetosum (subass. nova), Plantago atrata and Leontodon montanus community. The distribution of these communities within the karst basins was found to be related to variations in topographic and geomorphological parameters, such as altitude, slope, soil availability and stoniness. All the new associations proposed in this paper belong to the suballiance Leontopodio – Elynenion and to the alliance Seslerion apenninae, both of which are endemic to the central Apennines. In order to compare the plant community types identified within the Majella massif to similar associations found in the rest of the Apennine chain, synoptic tables were constructed. Finally, a comparative phytogeographical analysis of the alpine belt vegetation of the Apennines, Dinarides, southern Balkans and eastern Alps is presented.     

Gap analysis comparing protected areas with potential natural vegetation in Tuscany (Italy) and a GIS procedure to bridge the gaps

The aim was to compare the protected area (PA) network in Tuscany, Italy, with the areas referenced to different types of potential natural vegetation (PNV), to determine whether they are adequately represented for plant and habitat conservation purposes. For PNV, an existing but slightly updated and modified Italian Vegetation Series (VS) map was used. Each VS type corresponds to vegetation complexes that live under homogeneous environmental conditions and can each be considered an ecological land unit at the working scale employed here. Using GIS techniques, the geographic layers of PAs and VS were processed with spatial intersection to extract and quantify the VS contained within the boundaries of PAs. As a minimum conservation goal, we used the widely accepted 10% target threshold. It was found that, even though Tuscan PAs covered almost 20% of the total land surface, 94% of VS types resulted to be included in PAs with a percentage of at least 10% of their total area. The survey shows that the VS with the higher degree of inclusion in PAs are distributed in the Mediterranean Tuscany (coast and Tuscan Archipelago) and in some inner areas such as Apuan region, northern Apennines, Amiata Mt. and Farma-Merse Valley. Two VS types must be considered under-protected (i.e. contained in an existing PA network with percentages < 10%). We propose a simple GIS procedure based on certain priority assumptions: (a) existing PAs should be enlarged rather than new ones created and (b) their naturalness taken into account. This procedure produces a suitability map useful for identifying the best areas in which a local administration might look for solutions to bridge the gaps.     

Vegetation of rock crevices of the montane and colline zones in the Pamir-Alai and Tian Shan Mts in Tajikistan (Middle Asia)

The paper presents the results of phytosociological research conducted on the vegetation of rock habitats in the montane and colline zones of the Pamir-Alai Mountains (Tajikistan). During field surveys done in 2010–2013, 122 phytosociological relevés were sampled. Plant species were recorded according to the Braun-Blanquet cover-abundance scale. The vegetation studies were carried out generally in the western Pamir-Alai and south-western Tian Shan Mts, especially in the Zeravshan, Hissar, Hazratishoh, Darvaz, Rushan, Turkestan, Kuramin and Vanch ranges. A synopsis of the rock communities of the montane and colline zones in Tajikistan is proposed. The collected material presents most of the variability among the plant communities of rock crevices and fissures on calcareous as well as acidic substrates in the montane and colline zones in Tajikistan. As a result of field research and numerical analyses, eight associations have been distinguished: Scutellarietum hissaricae, Scutellarietumschugnanicae, Scutellarietum zaprjagaevii, Scutellarietum baldshuanicae, Tylospermetumlignosae, Dionysietum involucratae A. Nowak, M. Nobis, S. Nowak & A. Nobis, 2014, Nanorrhinetum ramosissimi, Campanuletum albertii and the community of Scutellaria adenostegia within new alliance – Caricion koshevnikovii. The main factors determining the species composition of classified associations seem to be the geography of the distribution range, elevation above sea level, rock type and exposition.     

Inclusion of phytosociological data in an index of vegetation fire danger: application and mapping on the Karst area around Trieste (Italy)

In this paper we propose a method to produce maps of fire danger index (FDI). The index includes in the formula a vegetation pyrogenic potential index (VPPI) based on floristic phytosociological data. The map of FDI is produced by integrating available phytosociological maps of vegetation, geomorphology and climate, using the Geographic Information System technology. The method is applied to an area of the coastal Classical Karst (NE-Italy). While fire risk maps based on the incidence of previous fires are useful for facilitating emergency operations, e.g. the allocation of fire fighting resources, we conclude that the proposed index offers an effective tool to plan actions for fire prevention.     

Korean traditional village forest (Ma-Eul-Soop) and potential natural vegetation: A case study on the Sachon-Ri Garo-Soop in Gyeongsangbuk-do,South Korea

Prediction of potential natural vegetation (PNV) requires autochthonous species distributions of a benchmark phytocoenosen. In Korea there remained unique forests or groves, i.e. a traditional Ma-Eul-Soop (MES), in the foothill zone which has influenced by the most aggressively anthropogenic activities since the Neolithic Age. Sachon-ri’s Garo-Soop (SGS) located at the southern-eastern Korean Peninsula has been reputed as the best conserved MES in Korea. We studied SGS for identifying components of the PNV. A total of 1,959 trees in the SGS were measured finescaled GPS locations and diameter at breast height (DBH). Distribution patterns of major species were interpreted by using the MES species ranking class (MSRC) framed by a matrix of phytogeographical and phytosociological autochthons. We consequently found that the SGS was composed of 29 autochthonous species. Cornus walteri, Hemiptelea davidii, Quercus aliena, Zelkova serrata, and Celtis sinensis were identified as the best matched PNV species to the floodplain behind riverbank. A lot of saplings occurring at the forest floor were the major components of PNV too. By the way many allochthonous species have been consistently introduced into the SGS in terms of a man-made forest of most previous works. We suggested that such PNV-unfriendly managements should be promptly stopped.     

Synergistic attraction of pine sawyer Monochamus saltuarius (Coleoptera: Cerambycidae) to monochamol and α‐pinene

Monochamus (Coleoptera: Cerambycidae) species are longhorn pine sawyers that serve as insect vectors of the pinewood nematode Bursaphelenchus xylophilus (Nematoda: Parasitaphelenchidae), which are responsible for debilitating pine wilt disease. An aggregation pheromone, 2‐(1‐undecyloxy)‐1‐ethanol (hereafter referred to as monochamol), was shown to be effective at attracting Monochamus species. However, attraction of the pine sawyers to aggregation pheromones varied depending on semiochemicals, including host plant volatiles and kairomones. In this study, we investigated the abilities of monochamol and the host‐plant volatiles α‐pinene and ethanol to attract M. saltuarius in a pine forest in Cheongsong, Gyeongsangbuk‐do, Korea. A total of 91 M. saltuarius (28 males and 63 females) were captured. The combination of monochamol (700 mg) with α‐pinene and ethanol exhibited a synergistic effect on attracting M. saltuarius (11.0 beetles per trap), whereas monochamol alone and a mixture of α‐pinene and ethanol resulted in the capture of 3.2 beetles and 3.6 beetles per trap, respectively. Our results suggest that multi‐funnel traps baited with a blend of monochamol, α‐pinene and ethanol are highly effective for monitoring M. saltuarius and M. alternatus in pine forests.     

Twenty‐five years of plant community dynamics and invasion in New Zealand tussock grasslands

Understanding how plant communities respond to plant invasions is important both for understanding community structure and for predicting future ecosystem change. In a system undergoing intense plant invasion for 25 years, we investigated patterns of community change at a regional scale. Specifically, we sought to quantify how tussock grassland plant community structure had changed and whether changes were related to increases in plant invasion. Frequency data for all vascular plants were recorded on 124, permanent transects in tussock grasslands across the lower eastern South Island of New Zealand measured three times over a period of 25 years. Multivariate analyses of species richness were used to describe spatial and temporal patterns in the vegetation. Linear mixed‐effects models were used to relate temporal changes in community structure to the level and rate of invasion of three dominant invasive species in the genus Hieracium while accounting for relationships with other biotic and abiotic variables. There was a strong compositional gradient from exotic‐ to native‐dominated plant communities that correlated with increasing elevation. Over the 25 years, small‐scale species richness significantly decreased and then increased again; however, these changes differed in different plant communities. Exotic species frequency consistently increased on some transects and consistently declined on others. Species richness changes were correlated with the level of Hieracium invasion and abiotic factors, although the relationship with Hieracium changed from negative to positive over time. Compositional changes were not related to measured predictors. Our results suggest that observed broad‐scale fluctuations in species richness and community composition dynamics were not driven by Hieracium invasion. Given the relatively minor changes in community composition over time, we conclude that there is no evidence for widespread degradation of these grasslands over the last 25 years. However, because of continuing weed invasion, particularly at lower elevations, impacts may emerge in the longer term.     

6. Peat-bog Begbunar (Osogovo Mountains,south-west Bulgaria): Four millennia of vegetation history

The aim of this study was to estimate the polleniferous potential of the most common plant communities and crops across the landscape through the study of pollen loads collected by Apis mellifera. We analysed pollen load samples from hives located in Mallorca (western Mediterranean) for one year and compared the results with a phytosociological study of the foraging area. We also analysed meteorological data to evaluate the influence of some environmental factors on the pollen production phenology. We identified 74 different pollen types (66.2% from native flora, 25.7% from exotic flora, and 8.1% from crops) belonging to 33 families. The highest polleniferous potentials in the landscape were attained by sclerophyllous forests (Quercetea ilicis) in spring, crops in autumn, wild annual rangelands (Stellarietea mediae) in winter and gardens in late summer. Our results suggest that the higher the coverage present in a vegetation class, the higher the quantity and diversity of pollen that may be provided to pollinators; therefore, its polleniferous potential may be higher. Some meteorological conditions (rainfall, temperature and daily hours of sunlight) may affect this potential through their influence on flowering phenology.     

Stipa zeravshanica sp. nov. (Poaceae), an endemic species from rocky walls of the western Pamir Alai Mountains (middle Asia)

Stipa zeravshanica M. Nobis sp. nov. from the western Pamir Alai Mts (Tajikistan) is described and illustrated. Morphologically, the new species is similar to S. gracilis, but it is easily distinguishable by pilose (vs. glabrous) lower part of the awn, and generally somewhat longer ligules of the vegetative shoots. Epidermal patterns of the lemma in S. zeravshanica and three other closely related taxa, namely S. gracilis, S. himalaica and S. orientalis, were examined by means of SEM. The main macro‐morphological and micro‐morphological characteristics differentiating all mentioned species are presented. Additionally, a lectotype for S. gracilis is designated. Based on phytosociological relevés prepared during field investigation in the western Pamir Alai Mts, a new plant association Asperulo albiflorae–Stipetum zeravshanicae, occurring on shelves and rocky crevices is also described.     

sRNA‐FISH: versatile fluorescent in situ detection of small RNAs in plants

Localization of mRNA and small RNAs (sRNAs) is important for understanding their function. Fluorescent in situ hybridization (FISH) has been used extensively in animal systems to study the localization and expression of sRNAs. However, current methods for fluorescent in situ detection of sRNA in plant tissues are less developed. Here we report a protocol (sRNA‐FISH) for efficient fluorescent detection of sRNAs in plants. This protocol is suitable for application in diverse plant species and tissue types. The use of locked nucleic acid probes and antibodies conjugated with different fluorophores allows the detection of two sRNAs in the same sample. Using this method, we have successfully detected the co‐localization of miR2275 and a 24‐nucleotide phased small interfering RNA in maize anther tapetal and archesporial cells. We describe how to overcome the common problem of the wide range of autofluorescence in embedded plant tissue using linear spectral unmixing on a laser scanning confocal microscope. For highly autofluorescent samples, we show that multi‐photon fluorescence excitation microscopy can be used to separate the target sRNA‐FISH signal from background autofluorescence. In contrast to colorimetric in situ hybridization, sRNA‐FISH signals can be imaged using super‐resolution microscopy to examine the subcellular localization of sRNAs. We detected maize miR2275 by super‐resolution structured illumination microscopy and direct stochastic optical reconstruction microscopy. In this study, we describe how we overcame the challenges of adapting FISH for imaging in plant tissue and provide a step‐by‐step sRNA‐FISH protocol for studying sRNAs at the cellular and even subcellular level.