ArticlePublisher preview available

Patterns of distribution, abundance and composition of forest terrestrial orchids

Authors:
Vladan Djordjević at University of Belgrade
Vladan Djordjević
Spyros Tsiftsis at Democritus University of Thrace
Spyros Tsiftsis
Dmitar Lakusic at University of Belgrade
Dmitar Lakusic
Slobodan Jovanović at University of Belgrade
Slobodan Jovanović
Show all 6 authorsHide
Read publisher preview
Download citation
To read the full-text of this research, you can request a copy directly from the authors.

Abstract and Figures

Many terrestrial orchid species are known to grow in forest ecosystems, but the patterns of their diversity and habitat specialization have not been sufficiently explored. Niche parameters and factors influencing the distribution, abundance and composition of forest orchids in the central Balkans were investigated in the present study. Outlying mean index (OMI) analysis was used to explore ecological niches of orchids and environmental factors affecting the patterns of their distribution. Indicator species analysis was performed to identify species presenting strong affinity for specific forest and bedrock types. In addition, similarity profile analysis was applied to classify orchid taxa into meaningful groups, whereas the multi-response permutation procedure was used to analyse differences of orchid composition between forest types. Out of a total of 40 orchid species and subspecies analysed, 29 showed significant niche marginality. The first three axes of the OMI analysis explained 68.13% of total variability. Light, temperature, moisture, nitrogen and altitude were found to be the factors most effectively influencing the distribution and abundance of orchids. Seven orchids had statistically significant strong affinity for specific forest types, whereas eight orchids were statistically significantly correlated with specific bedrock types. Overall, six ecological groups of orchids were distinguished. The results suggest that specialist orchids occur at the extreme ends of the light, temperature, soil pH, moisture and altitude gradients. Moreover, most specialist orchid species were found from low to middle elevations. The results provide a useful basis for the successful design of strategies for the conservation of forest orchids.
Figures - available from: Biodiversity and Conservation
This content is subject to copyright. Terms and conditions apply.
A preview of this full-text is provided by Springer Nature.
Learn more
Content available from Biodiversity and Conservation
This content is subject to copyright. Terms and conditions apply.
ORIGINAL PAPER
Patterns of distribution, abundance and composition
of forest terrestrial orchids
Vladan Djordjevic
´
1
•Spyros Tsiftsis
2,3
•Dmitar Lakus
ˇic
´
1
•Slobodan Jovanovic
´
1
•
Ksenija Jakovljevic
´
1
•Vladimir Stevanovic
´
1,4
Received: 10 April 2020 / Revised: 3 October 2020 / Accepted: 9 October 2020 /
Published online: 14 October 2020
ÓSpringer Nature B.V. 2020
Abstract
Many terrestrial orchid species are known to grow in forest ecosystems, but the patterns of
their diversity and habitat specialization have not been sufficiently explored. Niche param-
eters and factors influencing the distribution, abundance and composition of forest orchids in
the central Balkans were investigated in the present study. Outlying mean index (OMI)
analysis was used to explore ecological niches of orchids and environmental factors affecting
the patterns of their distribution. Indicator species analysis was performed to identify species
presenting strong affinity for specific forest and bedrock types. In addition, similarity profile
analysis was applied to classify orchid taxa into meaningful groups, whereas the multi-
response permutation procedure was used to analyse differences of orchid composition
between forest types. Out of a total of 40 orchid species and subspecies analysed, 29 showed
significant niche marginality. The first three axes of the OMI analysis explained 68.13% of
total variability. Light, temperature, moisture, nitrogen and altitude were found to be the
factors most effectively influencing the distribution and abundance of orchids. Seven orchids
had statistically significant strong affinity for specific forest types, whereas eight orchids
were statistically significantly correlated with specific bedrock types. Overall, six ecological
groups of orchids were distinguished. The results suggest that specialist orchids occur at the
extreme ends of the light, temperature, soil pH, moisture and altitude gradients. Moreover,
most specialist orchid species were found from low to middle elevations. The results provide
a useful basis for the successful design of strategies for the conservation of forest orchids.
Keywords Conservation Environmental factors Forest ecosystems Habitat specialism
Niche analysis Orchidaceae
Communicated by Daniel Sanchez Mata.
This article belongs to the Topical Collection: Forest and plantation biodiversity
Electronic supplementary material The online version of this article (https://doi.org/10.1007/s10531-020-
02067-6) contains supplementary material, which is available to authorized users.
&Vladan Djordjevic
´
vdjordjevic@bio.bg.ac.rs
Extended author information available on the last page of the article
123
Biodiversity and Conservation (2020) 29:4111–4134
https://doi.org/10.1007/s10531-020-02067-6(0123456789().,-volV)(0123456789().,-volV)
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
... The relationships between forests and orchids have increasingly drawn the attention of researchers, who have explored various ecological aspects of these interactions [16][17][18][19][20]. In the case of temperate terrestrial forest orchids, their growth is known to depend not only on climatic factors and edaphic factors, but also on biotic interactions [21][22][23]. These species depend on mutualistic mycorrhizal associations with fungi to absorb carbon from nearby trees, a process that is essential for their survival [24]. ...
... In the forest ecosystem, other factors can be crucial. Soil characteristics play a fundamental role in influencing the abundance and distribution of Orchidaceae [3,23]. The type of geological substrate is important at both regional and local levels. ...
... For soil aspects, although partly comparable among the study sites, local differences in lithology and soil properties may additionally affect the presence of orchids either directly (e.g., pH, Cation Exchange Capacity, nutrients, etc.) [23] or indirectly (e.g., tree height, forest composition) [3]. However, we acknowledge that the lack of enough replications in our study does not allow for fully disentangling the relative effects of tree community vs. soil features on the presence of orchids. ...
Naturalness and Tree Composition Determine the Abundance of Rare and Threatened Orchids in Mature and Old-Growth Abies alba Forests in the Northern Apennines (Italy)
Article
Full-text available
  • Mar 2025
Forest Orchidaceae are important for European temperate forests, yet their distribution and abundance have so far interested limited research. In three pure or mixed silver fir stands in the Foreste Casentinesi National Park (NP) (Northern Apennines, Italy) we analysed how structural traits in mature and old-growth forests affected orchid communities in terms of abundance of the main genera, trophic strategy and rarity in the NP. We established three 20 × 60 m plots to quantify the structure of living and dead tree community, including a set of old-growth attributes connected to large trees, deadwood, and established regeneration. In each plot, we measured the abundance of all orchid species and explored their behaviour according to the trophic strategy (autotrophy/mixotrophy, obligate mycoheterotrophy), rarity within the NP, and threatened status according to the IUCN Red List. We used multivariate ordination and classification techniques to assess plot similarities according to forest structure and Orchid Community and identify the main structural factors related to orchid features. The main structural factors were used as predictors of community traits. Forest composition (i.e., the dominance/abundance of silver fir) affected the presence of the main orchid genera: Epipactis were abundant in silver fir-dominated forests, Cephalanthera in mixed beech and fir forests. Interestingly, Cephalanthera could become limited even in beech-dominated conditions if fir regeneration was abundant and established. Old-growth attributes like the density of deadwood and large tree volume were important determinants of the presence of rare and mycoheterotrophic species. Our results provided a first quantitative description of forest reference conditions to be used in the protection and restoration of threatened and rare orchid species.
View
... The diversity patterns of orchids in large geographical areas are primarily determined by the evolutionary and migratory history, macroclimatic factors and the latitude and size of the area [8,[10][11][12][13][14][15][16], while at the regional level, the distribution of orchids is influenced by the physico-chemical properties of the soil, geological substrates, light regime, meso-and microclimates, altitude, habitat types and biotic factors, i.e., the specificity of mycorrhizal fungi and pollinators as well as the disturbance regime [17][18][19][20][21][22][23][24][25]. However, the importance of these factors varies from region to region, and there is no universal diversity pattern that applies to all orchids and regions. ...
... Although the Balkan Peninsula is recognised as an important centre of orchid diversity, the variation in orchid species richness at a spatial scale of 10 km grid cells and the influence of factors on patterns of orchid diversity have not been studied in the Central Balkans. In this region, patterns of orchid abundance have been studied, which refers to the variation in the population size of orchids, especially in grassland and herbaceous wetlands [21,46] and forest ecosystems [22]. Although the influence of environmental factors on orchid diversity patterns can be observed individually (e.g., climatic factors), in reality, there is a joint effect of several environmental factors [47]. ...
... Pinion nigrae forests also play an important role in determining the distribution of orchid species richness in the Central Balkans, which is explained by the significant presence of this type of vegetation, especially on serpentines, but also on the carbonate geological substrates of the study area. Due to high light intensity, this type of forest vegetation is often inhabited by orchids that prefer open grassland habitats and is therefore considered an alternative habitat for some typical grassland orchids [22]. ...
Factors Influencing Orchid Species Richness in the Central Balkans: The Importance of Belowground Organ Types
Article
Full-text available
  • Feb 2025
The Balkan Peninsula is considered one of the most important centres of orchid diversity in Europe. However, the patterns of orchid species richness in the Central Balkans have not been sufficiently studied so far. The aim of this study was, therefore, to identify the centres of orchid diversity and the factors that influence the spatial variation in orchid species richness in the Central Balkans. For the analyses, the area of the Central Balkans was divided into 10 × 10 km grid cells. The environmental variables determined for each grid cell and used in the analyses were altitude, bioclimatic variables, geological substrates and habitat types. A random forest (RF) analysis was used to identify the environmental predictors most strongly associated with species richness. In addition to the total number of taxa, orchids with three belowground organ types were analysed separately: (a) rhizomatous orchids, (b) orchids with palmately lobed and fusiform tubers ("palmate tuberous orchids") and (c) orchids with spherical or ovoid tubers ("ovoid tuberous orchids"). In the Central Balkans, 54 orchid species and subspecies have been recorded, and the most important centres of diversity are the Tara, Zvijezda, Jadovnik and Zlatar Mountains and the Ovčar-Kablar Gorge. In general, two groups of grid cells with the largest number of orchid taxa, i.e., hotspots, stood out: (1) grid cells with a large altitudinal range and (2) grid cells occupied by gorges and ravines. The most important gradients influencing orchid species richness are specific habitat types and altitudinal ranges, while climatic factors and geological substrates are less important. The most important factors affecting the richness of total and rhizomatous orchids are altitudinal range and habitat types (Abieti-Fagenion, Ostryo-Carpinion orientalis and Pinion nigrae forests), highlighting the important role of habitat heterogeneity. The maximum altitude, percentage of Abieti-Fagenion and Vaccinio-Picetea forests and the minimum value of the mean temperature of the driest quarter are the most important factors for determining the richness of palmate tuberous orchids, whereas the percentage of xero-thermophilous habitat types (Ostryo-Carpinion orientalis, Asplenietea trichomanis and Pinion nigrae) has the greatest influence on the richness of ovoid tuberous orchids. These results confirm the hypothesis concerning the origin and development of underground organs in orchids, emphasising that palmate tuberous orchids are best adapted to cold and humid habitat conditions, whereas ovoid tuberous orchids have the ability to grow in habitats with very warm and dry conditions. This study provides a good basis for better orchid conservation planning and underlines the importance of belowground strategies as a feature of orchid life history that should be considered when studying patterns of orchid diversity.
View
... Indeed, there is evidence that, in the Mediterranean bioclimate, calcareous and serpentine substrates are common favorable conditions for orchids in Sardinia in the Iglesiente mine district [67] and Barbagia [68], as well as elsewhere like in the Balkans [69]. As far as the metallophyte character of the investigated flora is concerned, the results showed a high presence (49%) of taxa with this characteristic. ...
The First Inventory of Sardinian Mining Vascular Flora
Article
Full-text available
  • Apr 2025
Citation: Boi, M.E.; Sarigu, M.; Fois, M.; Casti, M.; Bacchetta, G. The First Inventory of Sardinian Mining Vascular Flora. Plants 2025, 14, 1225. Abstract: Mining activities and associated waste materials pose significant environmental challenges, including soil, water, and air contamination, along with health risks to nearby populations. Despite the harsh conditions of metal-enriched soils and nutrient-poor sub-strates, certain plants known as metallophytes thrive in these environments. This study examined the vascular flora of Sardinia's abandoned mining sites, with a focus on identifying metallophytes and their potential role in phytoremediation. A comprehensive floristic checklist was compiled using literature, field surveys, and herbarium samples. Of the 652 taxa identified, 49% were metallophytes, with the majority categorized as facultative species. Notably, 27% of metallophytes were identified as suitable for phytostabilization, while 20% showed potential for phytoextraction. This study also highlighted the presence of endemic and endangered species, emphasizing the need for conservation efforts. The findings suggest that native metallophytes could play a key role in the ecological restoration of mining sites, though careful consideration of invasive species is necessary to avoid ecological disruption. This research provides valuable insights into the biodiversity of Sardinian mining sites and the potential for sustainable remediation strategies using native plants.
View
... The species is known as a generalist which grows on different bedrock types, across a wide altitudinal range and in diverse habitats, including different types of forest vegetation (Djordjević et al. 2020). However, three newly recorded populations of this species were specifically found in beech forests, while the population on Mt. ...
New records and noteworthy data of plants, algae and fungi in SE Europe and adjacent regions, 21
Article
Full-text available
  • Apr 2025
This paper presents new records and noteworthy data on the following taxa in SE Europe and adjacent regions: lichenised fungus Solorina bispora var. subspongiosa, saprotrophic fungi Geastrum berkeleyi and Marasmius epiphylloides, liverworts Bazzania trilo
View
... Cymbidium macrorhizon Lindl., a leafless terrestrial orchid in the genus Cymbidium, is native to tropical and subtropical Asia and northern Australia, and inhabits dense evergreen or deciduous broadleaf forests, as well as mixed pine forests (National Institute of Biological Resources 2021; Ogura-Tsujita et al. 2012;Suetsugu et al. 2018;Thakur and Dutt 2021). Especially, terrestrial orchid distribution and population formation are influenced by various factors, including topography, vegetation, mycorrhizal viability, soil physicochemical properties, such as organic matter, pH, and nutrients, and climate (Djordjević et al. 2020;Swarts and Dixon 2009). ...
View
... These plants possess specialized mycorrhizal communities and unique pollination mechanisms 5 , rendering them extremely sensitive to habitat changes 6,7 . Furthermore, orchids hold significant ornamental value and medicinal properties 8 , establishing them as "flagship" species within their ecological contexts 9 . Currently, wild orchids are under significant pressure to survive due to severe anthropogenic activities and the effects of global climate change. ...
Fungal community characteristics of the last remaining habitat of three paphiopedilum species in China
Article
Full-text available
  • Oct 2024
Paphiopedilum armeniacum, Paphiopedilum wenshanense and Paphiopedilum emersonii are critically endangered wild orchids. Their populations are under severe threat, with a dramatic decline in the number of their natural distribution sites. Ex situ conservation and artificial breeding are the keys to maintaining the population to ensure the success of ex situ conservation and field return in the future. The habitat characteristics and soil nutrient information of the last remaining wild distribution sites of the three species were studied. ITS high-throughput sequencing was used to reveal the composition and structure of the soil fungal community, analyze its diversity and functional characteristics, and reveal its relationship with soil nutrients. The three species preferred to grow on low-lying, ventilated and shaded declivities with good water drainage. There were significant differences in soil alkali-hydrolyzed nitrogen and available phosphorus among the three species. There were 336 fungal species detected in the samples. On average, there were different dominant groups in the soil fungal communities of the three species. The functional groups of soil fungi within their habitats were dominated by saprophytic fungi and ectomycorrhizae, with significant differences in diversity and structure. The co-occurrence network of habitat soil fungi was mainly positive. Soil pH significantly affected soil fungal diversity within their habitats of the three paphiopedilum species. The study confirmed that the dominant groups of soil fungi were significantly correlated with soil nutrients. The three species exhibit comparable habitat inclinations, yet they display substantial variations in the composition, structure, and diversity of soil fungi. The fungal functional group is characterized by a rich presence of saprophytic fungi, a proliferation of ectomycorrhizae, and a modest occurrence of orchid mycorrhizae. The symbiotic interactions among the soil fungi associated with these three species are well-coordinated, enhancing their resilience against challenging environmental conditions. There is a significant correlation between soil environmental factors and the composition of soil fungal communities, with pH emerging as a pivotal factor regulating fungal diversity. Our research into the habitat traits and soil fungal ecosystems of the three wild Paphiopedilum species has established a cornerstone for prospective ex situ conservation measures and the eventual reestablishment of these species in their native landscapes.
View
... Previous studies have extensively explored temperate European orchids, analysing the biotic and abiotic factors that influence their presence, abundance and distribution [4,55,[70][71][72], including species commonly grown in forest environments [73][74][75]. However, it is important to note that only a limited number of studies have been conducted to assess the potential future distribution of a few species in relation to climate change using a species distribution modelling approach [25,[76][77][78][79]. ...
Forest Orchids under Future Climate Scenarios: Habitat Suitability Modelling to Inform Conservation Strategies
Article
Full-text available
  • Jun 2024
Orchidaceae is one of the largest and most diverse families of flowering plants in the world but also one of the most threatened. Climate change is a global driver of plant distribution and may be the cause of their disappearance in some regions. Forest orchids are associated with specific biotic and abiotic environmental factors, that influence their local presence/absence. Changes in these conditions can lead to significant differences in species distribution. We studied three forest orchids belonging to different genera (Cephalanthera, Epipactis and Limodorum) for their potential current and future distribution in a protected area (PA) of the Northern Apennines. A Habitat Suitability Model was constructed for each species based on presence-only data and the Maximum Entropy algorithm (MaxEnt) was used for the modelling. Climatic, edaphic, topographic, anthropogenic and land cover variables were used as environmental predictors and processed in the model. The aim is to identify the environmental factors that most influence the current species distribution and the areas that are likely to contain habitats suitable for providing refuge for forest orchids and ensuring their survival under future scenarios. This will allow PA authorities to decide whether to invest more resources in conserving areas that are potential refuges for threatened species.
View
... The orchids can find enough light, cool temperatures, moisture, and other micronutrients they need in the forest to survive. These factors, together with elevation and the presence of soil nitrogen, are the primary determinants of the distribution and abundance of orchids, according to Djordjević et al. (2020). Some orchid species become habitat specialists due to these ecological factors (Kirillova et al. 2022). ...
Orchid diversity across different forest types on Mt. Malindang, Philippines
Article
Full-text available
  • Feb 2024
Labajo-Villantes Y, Cootes J, Luo Y-B, Nuneza OM. 2024. Orchid diversity across different forest types on Mt. Malindang, Philippines. Biodiversitas 25: 605-615. Mt. Malindang Range Natural Park is one of the most significant biodiversity and conservation areas and is considered a biodiversity hotspot. There have been many biodiversity studies on Mt. Malindang, but information on orchid taxonomy and distribution is limited. This study aimed to conduct a comprehensive inventory of orchids using an exploratory method. Samplings were conducted in different forest types: montane, mossy, dipterocarp, mixed dipterocarp, and almaciga. A total of 114 orchid species with 55 endemics were recorded. Three of these (Phalaenopsis x intermedia Lindl., Paphiopedilum hennisianum (M.W.Wood) Fowlie, and Paphiopedilum haynaldianum (Rchb.f.) Stein) were under Appendix I (can only be exported under specific circumstances). At the same time, 78 were under the Appendix II (follow controlled trades) categories of the Convention on International Trade in Endangered Species of Wild Fauna and Flora. P. hennisianum and P. haynaldianum, listed as endangered species globally, were observed in montane and mossy forests, respectively. Among the six sampling sites, the montane forest was the most species-rich, with the most endemics, followed by the mossy forest. Almaciga and mixed dipterocarp forests were closely related with 100% similarity. The results of this study are valuable to Mt. Malindang's ecological profile for more effective protection and conservation plans.
View
Soil Properties and Rhizosphere Microbes Community Structure Reveal Nitrogen Uptake Preferences and Nitrogen Use Efficiency of Two Ecotypes of Paphiopedilum micranthum
Article
Full-text available
  • Oct 2024
Paphiopedilum micranthum, an IUCN Red List species, is discontinuously distributed in the karst limestone mountain of southwest China and exhibits ecological specialization, typically through lithophytic and terrestrial ecotypes. Whether the distribution of rhizosphere bacteria and fungi in these different habitats is random or reflects soil preferences requires further investigation. A total of 73 samples from the core distribution areas in China, representing all habitats in two sites, were analyzed for soil differences by comparing edaphic properties and microbial community structure based on high-throughput sequencing of bacterial 16S rRNA genes and fungal ITS region sequences, alongside soil physiochemical data. The results showed no significant differences in microbial community richness and diversity across the heterogeneous habitats. However, significant differences in taxa were observed across various habitats. Dominant bacterial phyla included Actinobacteriota, Proteobacteria and Acidobacteriota, with dominant genera such as Crossiella, Pseudonocardia, 67-14, Mycobacterium and RB41. The primary fungal phyla were Basidiomycota and Ascomycota, featuring prominent genera such as Phlegmacium, Archaeorhizomyces, Trechispora, and Lepiota. There were 16 bacterial genera and 13 fungal genera associated with nitrogen transformation and fixation. Alkali-hydrolyzed nitrogen (AN) was identified as a main driver of soil bacterial and fungal community variation. Based on an analysis of soil physicochemical properties, ammonium nitrogen content was consistently higher than nitrate nitrogen across different habitats. Furthermore, across all heterogeneous habitats, P. micranthum showed no significant differences in nitrate nitrogen, ammonium nitrogen, or their ratio. The nitrogen-use efficiency of P. micranthum ranged from 7.73% to 9.87%, with the highest efficiency observed in the terrestrial habitat of Shedu. These results suggest that P. micranthum prefers habitats rich in organic matter and nitrogen, showing a preference for ammonium nitrogen uptake in natural conditions. Heterogeneous habitats affect plant nitrogen-use efficiency as well as changes in microbial community composition.
View
Fungal community characteristics of the last remaining habitat of three paphiopedilum species in China
Preprint
Full-text available
  • Apr 2024
Background: Paphiopedilum armeniacum, Paphiopedilum wenshanense and Paphiopedilum emersonii are critically endangered wild orchids. The population is threatened, and the number of natural distribution sites has plummeted. Ex situ conservation and artificial breeding are the keys to maintaining the population to ensure the success of ex situ conservation and field return in the future. The habitat characteristics and soil nutrient information of the last remaining wild distribution sites of the three species were studied. ITS high-throughput sequencing was used to reveal the composition and structure of the soil fungal community, analyze its diversity and functional characteristics, and reveal its relationship with soil nutrients. Results: The three species preferred to grow on low-lying, ventilated and shaded negative terrain with good water drainage. There were significant differences in soil alkali-hydrolyzed nitrogen and available phosphorus among the three species. There were 336 fungal species detected in the samples. On average, there were different dominant groups in the soil fungal communities of the three species. The functional groups of soil fungi in the habitats were dominated by saprophytic fungi and ectomycorrhizae, with significant differences in diversity and structure. The co-occurrence network of habitat soil fungi was mainly positive. Soil pH significantly affected soil fungal diversity in the habitats of the three paphiopedilum species. The study and analysis confirmed that the dominant groups of soil fungi were significantly correlated with soil nutrients. Conclusion: The three species have similar habitat preferences, but there are significant differences in soil fungal composition, community structure and diversity. The functional group of fungi contains abundant saprophytic fungi, ectomycorrhizae, and a small amount of orchid mycorrhizae. The symbiotic relationships of the three species of soil fungi were harmonious, which was conducive to resistance to adverse environments. Soil environmental factors were significantly correlated with soil fungal communities, and pH significantly controlled fungal diversity. Our study on the habitat characteristics and soil fungal communities of the three wild paphiopedilum species laid a foundation for future ex situ conservation and field return work.
View
EUNIS Habitat Classification: Expert system, characteristic species combinations and distribution maps of European habitats
Article
Full-text available
  • Dec 2020
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 dataset. 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.
View
Pollination Mechanisms are Driving Orchid Distribution in Space
Article
Full-text available
  • Jan 2020
Understanding the abundance and distribution patterns of species at large spatial scales is one of the goals of biogeography and macroecology, as it helps researchers and authorities in designing conservation measures for endangered species. Orchids, one of the most endangered groups of plants, have a complicated system of pollination mechanisms. Their survival strongly depends on pollination success, which then determines their presence and distribution in space. Here we concentrate on how pollination mechanisms (presence/absence of nectar) are associated with orchid species density and mean niche breadth along an altitudinal gradient in six different phytogeographical regions in the Czech Republic. We found differences between these regions in terms of orchid species numbers and density. The trend (hump-shaped curve) in species density of nectarless and nectariferous orchids were very similar in all phytogeographical regions, peaking between 300–900 m. The trend strongly depends on habitat cover and pollinator availability. In general, the most specialist species of orchids were found from low to middle altitudes. The association of altitude with the richness of orchid flora is much stronger than that with the biogeography. Climate change is a factor that should not be neglected, as it may affect the presence/absence of many species in the future.
View
A biogeographical and ecological classification of orchids in Turkey
Article
Full-text available
  • Apr 2020
  • PHYTOCOENOLOGIA
Aims: With its diverse vascular flora Turkey is well-known as a bridge between the Mediterranean basin, the Caucasus, and the Irano-Turanian region. Orchids are an important component of this diversity with 191 taxa, of which 39 taxa are Turkish endemics. This study aims to group the country’s orchid taxa into different phytogeographical regions, natural vegetation types, and altitudinal zones. Study area: Turkey. Methods: Field surveys were carried out from 1988 to 2017. The sampling areas were selected with good prior knowledge of orchid-containing terrains and habitats. Distribution data were revised and presented in a 10 km UTM grid. Geospatial data were classified and processed by ArcGIS software. Results: The Mediterranean region (70 taxa, 17 endemics) is the richest for orchid taxa distributed in only one phytogeographical region, followed by the Euro-Siberian (13 taxa, 2 endemics) and the Irano-Turanian (7 taxa, all endemics) regions. Species richness per habitat varied from Mediterranean landscapes of “natural Pinus brutia forests” (105 taxa, 12 endemics), followed by landscapes with “Quercus frainetto forest” (59 taxa, 3 endemics), “Oak-hornbeam forest” (55 taxa, 3 endemics), “Beech-spruce forest” (38 taxa, 1 endemic), “Quercus brantii forest steppe (arid)” (38 taxa, 5 endemics) to “Subalpine Juniper forest steppe” (34 taxa, 7 endemics). Orchids were found mainly at elevations of 0 to 500 m (138 taxa) and 501 to 1,000 m (123 taxa). At elevations over 1,000 m, the number of orchid taxa decreases, but 21 taxa were found at over 2,000 m. Some orchid taxa were found to be restricted to a particular altitudinal zone and phytogeographical region. Conclusion: This study provides a general framework for relevant orchid conservation efforts in Turkey. Key habitats from different phytogeographical regions and altitudinal zones should be prioritised. For such areas, a multi-layered conservation planning approach from country to habitat level across Turkey needs to be developed. Complementary approaches should be promoted in neighbouring countries too.
View
Patterns of orchid species richness and composition in relation to geological substrates
Article
Full-text available
  • Dec 2019
  • WULFENIA
Terrestrial orchids in Europe are known for their great species richness that occurs on calcareous geological substrates. However, the knowledge of orchids that grow on other bedrock types is limited. In this paper, the richness and composition of orchid species in relation to geological substrates in the central Balkans (western Serbia) were investigated. For identification of floristic similarities among the geological substrates, the clustering method based on Jaccard distances and unweighted pair-group average hierarchical sorting strategy (UPGMA) was used. In addition, multi-response permutation procedure (MRPP) was applied to analyse the differences in orchid composition between bedrock types. Orchid species and subspecies were recorded on ten bedrock types. The greatest number of orchids was recorded on limestone-dolomite (52 species and subspecies), ophiolitic mélange (40) and schists-gneiss-phyllites (31). However, a significant number of orchid species and subspecies were recorded on carbonate clastites, ultramafics, andesite-dacite-porphyrite and Quaternary sediments. On the other hand, a smaller number of orchid species were recorded on acidic igneous rocks (quartz latite and granodiorite), indicating that they are less favourable for orchids. The cluster analysis of floristic similarities allowed the identification of seven sets of geological substrates. The greatest floristic similarity was identified between limestone-dolomite and ophiolitic mélange as well as schists-gneiss-phyllites and ultramafics. However, an analysis of the composition of orchid flora in terms of abundance and frequency of orchids indicated that carbonate substrates (limestone-dolomite and carbonate clastites) differ statistically from the highest number of bedrock types. Cephalanthera damasonium, C. longifolia, Dactylorhiza sambucina, Epipactis helleborine, Gymnadenia conopsea, Neottia nidus-avis and Platanthera bifolia were found to grow on nine bedrock types, suggesting their great ecological plasticity and adaptability. This study emphasizes the important role of geological substrates in the richness, composition and distribution of orchid species in the central Balkans, pointing out that many species, known to date to occur exclusively on calcareous substrates, also grow on different silicate substrates and ultramafics. The results indicate that differences in the species richness and composition of orchids which are related to the geological substrates of their habitats should be taken into account when designing orchid conservation and management strategies.
View
Identification of drivers of landscape distribution of forest orchids using germination experiment and species distribution models
Article
Full-text available
  • Jun 2019
  • OECOLOGIA
The family of orchids involves a number of critically endangered species. Understanding of drivers of their landscape distribution could provide a valuable insight into their decline. Our objectives were to develop models predicting distribution of selected orchid species—four co-occurring forest orchid species, Cephalanthera rubra, Epipactis atrorubens, E. helleborine, and Neottia nidus-avis—at a landscape scale using a wide range of habitat characteristics. Subsequently, we compared the model predictions with species occurrence and the results of the field germination experiment while considering two germination stages—asymbiotic (early stage) and symbiotic. And finally, we attempted to identify possible drivers of species’ landscape distribution (i.e., dispersal, availability of habitat patches, or fungal associates). We have discovered that different habitat characteristics determined the distribution of different orchids. The species also differed in terms of availability of suitable habitat patches and patch occupancy (the highest being E. atrorubens with 80%). Landscape distribution of the species was primarily restricted by the availability of fungal associates (the most important factor for C. rubra) and by the availability of suitable habitat patches (the most important in case of N. nidus-avis). Despite expected easy dispersal of spores, orchid distribution seems to be limited by the availability of fungal associates in the landscape. In contrast, the availability of orchid seeds does not seem to limit their distribution. These results can provide useful guidelines for conservation of the studied species.
View
Roadside verges and cemeteries: Comparative analysis of anthropogenic orchid habitats in the Eastern Mediterranean
Article
Full-text available
  • May 2019
Several important habitats have become threatened in the last few centuries in the Mediterranean Basin due to major changes adopted in land‐use practices. The consequent loss of natural and seminatural orchid habitats leads to the appreciation of small anthropogenic habitats, such as cemeteries and roadside verges. Colonization of cemeteries and roadside verges by orchids has long been known, but no study to date compared the suitability of these two anthropogenic habitats for orchids. Therefore, in this paper our aim was to survey cemeteries and roadside verges and to compare these two habitats regarding their role in conserving Mediterranean terrestrial orchids. We conducted field surveys in three Mediterranean islands, Cyprus, Crete, and Lesbos, where both cemeteries and roadside verges were sampled on a geographically representative scale. We found a total of almost 7,000 orchid individuals, belonging to 77 species in the two anthropogenic habitat types. Roadside verges hosted significantly more individuals than cemeteries in Crete and Lesbos, and significantly more species across all three islands. Our results suggest that although cemeteries have a great potential conservation value in other parts of the world, intensive maintenance practices that characterized cemeteries in these three islands renders them unable to sustain valuable plant communities. On the other hand, roadside verges play a prominent role in the conservation of Mediterranean orchids in Cyprus and Greece. The pioneer status of roadside verges facilitates their fast colonization, while roads serve as ecological corridors in fragmented landscapes.
View
Role of way of life, latitude, elevation and climate on the richness and distribution of orchid species
Article
Full-text available
  • Jan 2019
  • BIODIVERS CONSERV
We are experiencing climate warming that is likely to affect all regions worldwide, although in a different manner, when its consequences (e.g. increase of temperature, lower seasonality, lower environmental stochasticity) are considered. Thus, our prediction of how global change will affect distribution and survival of species can be estimated by using our knowledge, how species richness and distribution is related with latitude and elevation. We used 193 terrestrial orchid species and subspecies as an example and we classified them according to their root system. This trait represents the evolution of strategies for underground storage of resources and resource acquisition as well as a characteristics that can be described as a life history trait. Classification of orchid life history traits that focus on belowground strategies has never been examined in a macroecological study. We then explored the associations between species richness, mean niche breadth and mean distribution on one hand and selected predictors on the other hand, using regression techniques for all orchids, and then for their subsets with different root systems. The predictive power, as well as the significance of the predictors, was also tested using polynomial second order generalized linear models. Species richness for the three belowground strategies was significantly affected by the predictors, whereas their mean niche breadth and mean distribution were largely dependent on their evolutionary history. The correlations of mean niche breadth and mean distribution with maximum elevation, latitude and longitude were significant for all orchid taxa and their subsets. All the variables together accounted for almost 50% or more of the variance in each of the subsets. The maximum elevation was the most significant factor for rhizomatous and intermediate orchids, whereas minimum temperature in the coldest month was highly significant for the tuberous orchids. Spatial distribution of Greek orchids is associated with a combination of elevation, latitude and climate. The distributions of rhizomatous and intermediate orchids are mainly associated with the orographic configuration of Greece, whereas the tuberous orchids are widely distributed in the southern, central and north-western areas of Greece, where most of them are limited by the harsh winter climate.
View
Effectiveness of corridors varies among phytosociological plant groups and dispersal syndromes
Article
Full-text available
In agricultural landscapes, semi-natural habitats are scarce and remaining habitat patches are largely isolated. However, linear landscape elements might facilitate dispersal of plant species through the agricultural landscape matrix. We investigated the following research questions: 1. are open linear landscape elements (LLE) effective corridors for dispersal of vascular plant species? 2. Which plant species, with respect to phytosociological group and dispersal-distance class, do use LLE as corridors? 3. To which extent is floristic similarity of communities influenced by dispersal through corridors? Field work was carried out in agricultural landscapes of Northwest Germany. We sampled 50 vegetation relevés on open linear landscape elements i.e. field margins (incl. road verges) and ditches, in eight 1-km² study areas. Then, we calculated Jaccard similarities of all plot pairs within study areas using either all species or only species of certain phytosociological groups or dispersal-distance classes. We assessed the isolation of the plots from each other using both Euclidean distance and resistance distance along LLE. Resistance distance reflected the degree of connectivity of the LLE network between the plots. A stronger effect on Jaccard similarity of resistance distance compared to Euclidean distance would indicate corridor dispersal of plants through LLE. Relationships between Jaccard similarity and the two isolation measures were analysed with Generalised Linear Mixed Models. Resistance distance of LLE had a stronger negative effect on Jaccard similarity than Euclidean distance in field margins, but not in ditches. This was found for species of ‘meadows and pastures’ and short to medium dispersal distance. In plot pairs that were highly connected by LLE, the models suggested that roughly 20% of all species occurred in both plots due to dispersal through LLE. Other species groups did not respond more strongly to resistance distance than to Euclidean distance. We conclude that linear landscape elements in agricultural landscapes are effective corridors for dispersal of plant species that are confined to semi-natural habitats, such as traditional grasslands, and lack mechanisms of long-distance dispersal.
View
Orchid seed sensitivity to nitrate reflects habitat preferences and soil nitrate content
Article
  • Sep 2019
Orchids are distributed around the world, however, the factors shaping their specific distribution and habitat preferences are largely unknown. Moreover, many orchids are at risk of becoming threatened as landscapes change, sometimes declining without apparent reason. One important factor affecting plant distribution is nutrient levels in the environment. Nitrates can inhibit not only orchid growth and persistence, but also seed germination. We used in vitro axenic cultures to exactly determine the germination sensitivity of seven orchid species to nitrates and correlated this with soil properties of the natural sites and with the species’ habitat preferences. We found high variation in response to nitrate between species. Orchids from oligotrophic habitats were highly sensitive, while orchids from more eutrophic habitats were almost insensitive. Sensitivity to nitrate was also associated with soil parameters that indicated a higher nitrification rate. Our results indicate that nitrate can affect orchid distribution via direct inhibition of seed germination. Nitrate levels in soils are increasing rapidly due to intensification of agricultural processes and concurrent soil pollution, and we propose this increase could cause a decline in some orchid species.
View
Neottia cordata (Orchidaceae) at its southernmost distribution border in Europe: Threat status and effectiveness of Natura 2000 Network for its conservation
Preprint
  • Apr 2019
  • J NAT CONSERV
Neottia cordata is an orchid species that exclusively occurs in Picea abies subsp. abies and Pinus sylvestris stands and forms its southernmost distribution limits in Greece. Over the last decade, it has been recorded in numerous sites, in which it forms small colonies, both in terms of population and area. This research re-evaluates the threat status of Neottia cordata based on updated chorological and population data; it also checks the effectiveness of the Natura 2000 Network as far as its conservation is concerned by applying the MaxEnt model. The recently collected data justify the classification of N. cordata as "Vulnerable". The most significant variables that affect its distribution were found to be the vegetation type and the precipitation of the warmest quarter. The results of the MaxEnt model suggest that: (i) the Natura 2000 Network is characterised by areas of higher habitat suitability values when compared to the areas that fall outside this network; and, (ii) Picea abies subsp. abies forests are more suitable for its conservation than those of Pinus sylvestris. It is concluded that the most appropriate management measure for the species conservation is the maintenance of the tree layer canopy closed.
View