Climatic drivers of dry grassland phylogenetic diversity in the Republic of Macedonia - Online supplement

Climatic gradients can be used to predict the extent to which climate drives biodiversity and to which biodiversity may be affected by global climate changes. Climate and evolutionary history are linked by the ecological adaptations of species and the history of Earth’s climate. If so, phylogenetic diversity may be a good metric to estimate biodiversity. We aimed to test whether the phylogenetic diversity of Macedonian dry grasslands was related to climatic variables. We sampled 575 plots, identifying the species and building a phylogenetic tree for them. We calculated two metrics of phylogenetic diversity and regressed them against climatic variables. We also tested whether there were nodes in the tree responsible for the main observed spatial patterns of phylogenetic diversity. We found a strong signature of evolutionary history in species sorting across a gradient driven by climate in Macedonian dry grasslands. First, the amount of evolutionary history decreased towards drier and more seasonal climates, suggesting a phylogenetic niche conservatism. Second, there was an air temperature filter and a temperature seasonality filter, acting in opposite directions and leading to phylogenetic clustering. Third, there were few nodes in the phylogenetic tree with high degrees of allopatry, associated with clades that differed not only in their geographic distribution, but also in their climatic preferences. Macedonian dry grassland communities developed over centuries of traditional land use but are threatened nowadays by human activities. The use of phylogenetic approaches may lead to more effective conservation policies and help us preserve this highly diverse vegetation

Relation between boundaries of protected areas and the distribution of vulnerable natural habitats – a case study from Sharri National Park, SE Europe

A growing threat to areas designed to protect habitats with high biodiversity has been noticed. In order to assess the present level of threat, the correlation between the factual situation of natural habitats and the boundary of protected area was studied in the massif of Luboten, Sharri NP. 45 phytosociological relevès were made in the studied site, all habitat types were recorded and notes on presence of rare and endemic plant taxa were taken. It was noticed that within the massif of Luboten, Sharri NP, an endangered natural habitat of subalpine moist tall herbs is not covered within the strictly protected area. The Moesian hogweed tall herb communities with Cirsium appendiculatum Griseb., as the most distinctive plant species, are known to harbor several endemic and rare plant species. To further add conservation importance, in these habitats with narrow distribution and fragile environment, there is one South-European Orophilous plant species (Willemetia stipitata), as well as 12 Balkan endemic plant taxa. The corresponding recorded plant association is Doronico gigantei-Cirsietum appendiculati Horv. ex Quez. Based on the obtained data on the situation of this habitat, we highly suggest extending the strictly protected area for 0.56 km2 into the NW direction of the western slope

Blysmo compressi-Eriophoretum latifoliae ass. nova, a new association of the Caricion fuscae alliance from the Sharri Mountains

The sedge-moss vegetation of the moderately to low calcium-rich slightly acidic fens of the Caricion fuscae alliance depends on a very specific combination of ecological and climatic conditions to thrive. Until recently, the classification of this vegetation group was complicated by its rarity on the southern edges of its range in Europe. As part of a larger database of phytocenological relevés carried out in Mt. Luboten, we came across an interesting group of 15 relevés on fen vegetation sites. We were curious to know if this plant community was a previously known association or if it might represent something new within this alliance. We compiled a separate dataset at JUICE that includes four plant communities from this alliance, along with our 15 original releves. The classification was based on modified TWINSPAN and beta-flexible clustering as a numerical classification method, with OPTIMCLASS determining the appropriate number of clusters. Five associations were clearly delineated, with the four associations taken from the literature sources clearly grouped individually and a new, fifth association appearing as separate, with completely unique characteristics. This new association: Blysmo compressi-Eriophoretum latifoliae occurs at elevations of ~ 1650 m a.s.l. on NE and NW slopes of the mountain. With this work we offer the description of a new high-mountain fen association. These associations may play an important syntaxonomic role as more Balkan data become available on this alliance. The sedge-moss and fen vegetation in the Balkans is particularly rare and characterised by a very diverse and specific vegetation, so it rightly deserves more attention from vegetation scientists and conservation authorities

Testing macroecological abundance patterns: The relationship between local abundance and range size, range position and climatic suitability among European vascular plants

Aim: A fundamental question in macroecology centres around understanding the relationship between species' local abundance and their distribution in geographical and climatic space (i.e. the multi‐dimensional climatic space or climatic niche). Here, we tested three macroecological hypotheses that link local abundance to the following range properties: (a) the abundance-range size relationship, (b) the abundance-range centre relationship and (c) the abundance-suitability relationship. Location: Europe. Taxon: Vascular plants. Methods: Distribution range maps were extracted from the Chorological Database Halle to derive information on the range and niche sizes of 517 European vascular plant species. To estimate local abundance, we assessed samples from 744,513 vegetation plots in the European Vegetation Archive, where local species' abundance is available as plant cover per plot. We then calculated the 'centrality', that is, the distance between the location of the abundance observation and each species' range centre in geographical and climatic space. The climatic suitability of plot locations was estimated using coarse‐grain species distribution models (SDMs). The relationships between centrality or climatic suitability with abundance was tested using linear models and quantile regression. We summarized the overall trend across species' regression slopes from linear models and quantile regression using a meta‐analytical approach. Results: We did not detect any positive relationships between a species' mean local abundance and the size of its geographical range or climatic niche. Contrasting yet significant correlations were detected between abundance and centrality or climatic suitability among species. Main conclusions: Our results do not provide unequivocal support for any of the relationships tested, demonstrating that determining properties of species' distributions at large grains and extents might be of limited use for predicting local abundance, including current SDM approaches. We conclude that environmental factors influencing individual performance and local abundance are likely to differ from those factors driving plant species' distribution at coarse resolution and broad geographical extents

Mapping species richness of plant families in European vegetation

Aims: Biodiversity is traditionally studied mostly at the species level, but biogeographical and macroecological studies at higher taxonomic levels can provide valuable insights into the evolutionary processes at large spatial scales. Our aim was to assess the representation of vascular plant families within different vegetation formations across Europe. Location: Europe. Methods: We used a data set of 816,005 vegetation plots from the European Vegetation Archive (EVA). For each plot, we calculated the relative species richness of each plant family as the number of species belonging to that family divided by the total number of species. We mapped the relative species richness, averaged across all plots in 50 km × 50 km grid cells, for each family and broad habitat groups: forests, grasslands, scrub and wetlands. We also calculated the absolute species richness and the Shannon diversity index for each family. Results: We produced 522 maps of mean relative species richness for a total of 152 vascular plant families occurring in forests, grasslands, scrub and wetlands. We found distinct spatial patterns for many combinations of families and habitat groups. The resulting series of 522 maps is freely available, both as images and GIS layers. Conclusions: The distinct spatial patterns revealed in the maps suggest that the relative species richness of plant families at the community level reflects the evolutionary history of individual families. We believe that the maps and associated data can inspire further biogeographical and macroecological studies and strengthen the ongoing integration of phylogenetic, functional and taxonomic diversity concepts.MV, IA, JPC, ZL, IK, AJ and MC were funded by the Czech Science Foundation, programme EXPRO (project no. 19-28491X); JDi by the Czech Science Foundation (18-02773S); IB and JAC by the Basque Government (IT936-16); AČ by the Slovenian Research Agency (ARRS, P1-0236); AK by the National Research Foundation of Ukraine (project no. 2020.01/0140); JŠ by the Slovak Research and Development Agency (APVV 16-0431); KV by the National Science Fund (Contract DCOST 01/7/19.10.2018)

EUNIS Habitat Classification: Expert system, characteristic species combinations and distribution maps of European habitats

Aim: The EUNIS Habitat Classification is a widely used reference framework for European habitat types (habitats), but it lacks formal definitions of individual habitats that would enable their unequivocal identification. Our goal was to develop a tool for assigning vegetation‐plot records to the habitats of the EUNIS system, use it to classify a European vegetation‐plot database, and compile statistically‐derived characteristic species combinations and distribution maps for these habitats. Location: Europe. Methods: We developed the classification expert system EUNIS‐ESy, which contains definitions of individual EUNIS habitats based on their species composition and geographic location. Each habitat was formally defined as a formula in a computer language combining algebraic and set‐theoretic concepts with formal logical operators. We applied this expert system to classify 1,261,373 vegetation plots from the European Vegetation Archive (EVA) and other databases. Then we determined diagnostic, constant and dominant species for each habitat by calculating species‐to‐habitat fidelity and constancy (occurrence frequency) in the classified data set. Finally, we mapped the plot locations for each habitat. Results: Formal definitions were developed for 199 habitats at Level 3 of the EUNIS hierarchy, including 25 coastal, 18 wetland, 55 grassland, 43 shrubland, 46 forest and 12 man‐made habitats. The expert system classified 1,125,121 vegetation plots to these habitat groups and 73,188 to other habitats, while 63,064 plots remained unclassified or were classified to more than one habitat. Data on each habitat were summarized in factsheets containing habitat description, distribution map, corresponding syntaxa and characteristic species combination. Conclusions: EUNIS habitats were characterized for the first time in terms of their species composition and distribution, based on a classification of a European database of vegetation plots using the newly developed electronic expert system EUNIS‐ESy. The data provided and the expert system have considerable potential for future use in European nature conservation planning, monitoring and assessment

Distribution maps of vegetation alliances in Europe

Aim: The first comprehensive checklist of European phytosociological alliances, orders and classes (EuroVegChecklist) was published by Mucina et al. (2016, Applied Vegetation Science, 19 (Suppl. 1), 3–264). However, this checklist did not contain detailed information on the distribution of individual vegetation types. Here we provide the first maps of all alliances in Europe. Location: Europe, Greenland, Canary Islands, Madeira, Azores, Cyprus and the Caucasus countries. Methods: We collected data on the occurrence of phytosociological alliances in European countries and regions from literature and vegetation-plot databases. We interpreted and complemented these data using the expert knowledge of an international team of vegetation scientists and matched all the previously reported alliance names and concepts with those of the EuroVegChecklist. We then mapped the occurrence of the EuroVegChecklist alliances in 82 territorial units corresponding to countries, large islands, archipelagos and peninsulas. We subdivided the mainland parts of large or biogeographically heterogeneous countries based on the European biogeographical regions. Specialized alliances of coastal habitats were mapped only for the coastal section of each territorial unit. Results: Distribution maps were prepared for 1,105 alliances of vascular-plant dominated vegetation reported in the EuroVegChecklist. For each territorial unit, three levels of occurrence probability were plotted on the maps: (a) verified occurrence; (b) uncertain occurrence; and (c) absence. The maps of individual alliances were complemented by summary maps of the number of alliances and the alliance–area relationship. Distribution data are also provided in a spreadsheet. Conclusions: The new map series represents the first attempt to characterize the distribution of all vegetation types at the alliance level across Europe. There are still many knowledge gaps, partly due to a lack of data for some regions and partly due to uncertainties in the definition of some alliances. The maps presented here provide a basis for future research aimed at filling these gaps

Early spring ephemeral therophytic non-nitrophilous grasslands as a habitat of various species of Romulea in the southern Balkans

The work deals with habitats of Romulea bulbocodium and Romulea linaresii ssp. graeca in the southern Balkans. Both species appear in early spring ephemeral therophytic non-nitrophilous grasslands in regions under the influence of the Mediterranean climate. These communities are classified within the Romulion alliance, which encompasses such communities from the eastern Mediterranean area. It was established that the main climatic factor causing the diversity of these communities is seasonality in precipitation and temperature. Two associations are presented, as Lagopo-Poetum bulbosae and Romuleo graecae-Poetum bulbosae

Suha travišča Erysimo-Trifolietum v severovzhodnem delu Republike Makedonije

The study deals with the distribution and classification of the association Erysimo-Trifolietum Micevski 1977 (alliance Trifolion cherleri Micevski 1971, order Astragalo-Potentilletalia Micevski 1971, class Festuco-Brometea Br. Bl. et Tx. 1943). The association develops on siliceous bedrock of the northern and northeastern parts of the Republic of Macedonia. After the research of dry grasslands in the region of Kratovo, enough information was available to prepare a synthetic overview of the association Erysimo-Trifolietum. Within the frame of this association, two new subassoaciations are described – subass. scleranthetosum subass. nova and subass. brachypodietosum subass. nova. Analysis of geoelements showed that sub-Mediterranean species are the most numerous and analysis of life forms provided evidence of their therophyto-hemicryptophytic physiognomy. The paper also presents the localities of occurrence, their floristic composition, synecological characteristics, life forms incidence and areal types.Raziskava se ukvarja z razširjenostjo in klasifikacijo asociacije Erysimo-Trifolietum Micevski 1977 (zveza Trifolion cherleri Micevski 1971, red Astragalo-Potentilletalia Micevski 1971, razred Festuco-Brometea Br. Bl. et Tx. 1943). Asociacija se pojavlja na silikatni podlagi v severnem in severovzhodnem delu Republike Makedonije. Po zaključenih raziskavah v okolici Kratova, smo razpolagali z zadostno količino podatkov, da smo lahko pripravili sintetični pregled asociacije Erysimo-Trifolietum. V okviru asociacije smo opisali dve novi subasociaciji – subasociacijo sclerathetosum subass. nova in brachypodietosum subass. nova. Analiza horološkega spektra je pokazala, da so v združbah najbolj pogoste submediteranske vrste, analiza življenjskih oblik pa kaže na terofitsko-hemikriptofitski značaj te asociacije. Delo prikazuje lokalitete, kjer se združbe pojavljajo, floristično zgradbo, sinekološke značilnosti, spekter življenjskih oblik in horološki spekter

Relationships between vegetation of Macedonian pine (Pinus peuce Griseb.) and different types of soils on which it develops

This paper deals with relationships between vegetation of Macedonian pine (Pinus peuce Griseb.) and soils developed on different parent materials on the territory of the Republic of North Macedonia. We analysed the floristic composition of its communities at localities on limestone, on scree of dolomite marble and on scree of silicate (glaciofluvial deposit). On limestone and scree of dolomite marble, rendzinas on hard limestone and dolomite have developed, and on silicate parent material brown forest soils – (cambisols). The vegetation was sampled according to the Braun-Blanquet approach. Detrended correspondence analysis (DCA) and Ellenberg’s indicator values were used for ecological interpretation of the vegetation patterns. The mechanical and chemical properties of soil and textural classes were also processed. An evident increased presence of carbonates in the soil of scree of dolomite marble on Nidže Mountain was observed, unlike that on Shar Mountain which has formed on typical limestone. Although it is a forest community dominated by the same species, differences between the massifs, the precipitation regime, geology, differences in soil properties in relation to the appearance of carbonates and pH values, and other factors, result in differences in their floristic composition and are the reason for the distinction between the two groups. On silicate terrain on Nidže Mountain, Macedonian pine forests have also developed on brown forest soils (cambisols), with a different floristic composition to that of the other group on different parent material - carbonate (dolomite and limestone)