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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. [ABSTRACT FROM AUTHOR]

This work presents a summary of cytogenetic data, including new information, on several species within the tribe Neottieae, with an update of the karyotype for 23 species belonging to the genera Cephalanthera, Limodorum, Epipactis, and Neottia (including Listera). Each of these four genera also presents distinctive chromosomal features, such as bimodal karyotypes. Our research includes insights into the distribution of constitutive heterochromatin, measured using C-banding and, in some cases, specific fluorochromes for the detection of A-T- and G-C-rich DNA. In the Epipactis group, it is noteworthy that when using the Giemsa banding technique, certain species (e.g., E. placentina, E. meridionalis) with a chromosome number of 2n = 38 were observed to exhibit a conspicuous wide band of constitutive heterochromatin on the long arm of the third pair in a subcentromeric position, resembling what has been observed in E. helleborine. These differences also have the potential to contribute to the diversification of these species. Based on the karyological results obtained, a hypothesis regarding the origin of certain species within the E. helleborine group is proposed. Additionally, karyological analyses conducted on a specimen of E. microphylla revealed chromosome counts ranging from 36 to 40. Somatic metaphases exhibited evident structural alterations in certain chromosomes, showing rearrangements probably caused by translocation phenomena. Based on the data obtained from the species within the studied genera, it is conceivable that variations in chromosomes, both structurally and in the distribution of constitutive heterochromatin, exert a significant influence on the evolution of the karyotype. Moreover, in many entities belonging to the Neottieae tribe, these processes may also contribute to the diversification of the phenotype in some instances. [ABSTRACT FROM AUTHOR]

SPME analysis of the scent of Epipactis microphylla showed the presence of limonene as the main component of the scent. Other components were 2,4,4,6,6,8,8-heptamethyl-1-nonene, pentadecane, and heptadecane. The scent of Epipactis palustris was characterized by pentadecane, 2,4,4,6,6,8,8-heptamethyl-1-nonene, and heptadecane. The scent of Neottia nidus avis showed the presence of kaur-16-ene as the main component of the scent. Other components were heinecosane, tetradecane, pentadecane, hexadecane, heptadecane, and 5,9,13-trimethyl-4,8,12-tetradecanal. The scent of Neottia ovata is due to pentadecane, hexadecane, and heptadecane. The scent of Limodorum abortivum showed the presence of 2,4,4,6,6,8,8-heptamethyl-1-nonene, pentadecane, hexadecane, heptadecane, and 2-(dodecyloxy)-ethanol. [ABSTRACT FROM AUTHOR]

Epipactis albensis is an obligate autogamous orchid species which is widely distributed in central European countries. However, its phytocoenological affinities are not well known. They were investigated in this study based on 17 phytosociological relevés from the Czech Republic, Slovakia, and Poland obtained from the available literature and database sources, where the species has its centre of occurrence. The cluster analyses revealed three groups of vegetation types belonging to the hardwood floodplain forests of the Fraxino-Quercion roboris alliance, the riparian alder forests of Alnion incanae and the beech forests of Fagion sylvaticae. Brachypodium sylvaticum, Circaea lutetiana, Geum urbanum, and Urtica dioica (frequency 76%), followed by Dactylis glomerata agg. and the alien plant Impatiens parviflora (65%) are the most co-occurring species. This phytosociological study has highlighted the need for further and more detailed research supported by the collection of new vegetation data from across the range of E. albensis.

This work presents a summary of cytogenetic data, including new information, on several species within the tribe Neottieae, with an update of the karyotype for 23 species belonging to the genera Cephalanthera, Limodorum, Epipactis, and Neottia (including Listera). Each of these four genera also presents distinctive chromosomal features, such as bimodal karyotypes. Our research includes insights into the distribution of constitutive heterochromatin, measured using C-banding and, in some cases, specific fluorochromes for the detection of A-T- and G-C-rich DNA. In the Epipactis group, it is noteworthy that when using the Giemsa banding technique, certain species (e.g., E. placentina, E. meridionalis) with a chromosome number of 2n = 38 were observed to exhibit a conspicuous wide band of constitutive heterochromatin on the long arm of the third pair in a subcentromeric position, resembling what has been observed in E. helleborine. These differences also have the potential to contribute to the diversification of these species. Based on the karyological results obtained, a hypothesis regarding the origin of certain species within the E. helleborine group is proposed. Additionally, karyological analyses conducted on a specimen of E. microphylla revealed chromosome counts ranging from 36 to 40. Somatic metaphases exhibited evident structural alterations in certain chromosomes, showing rearrangements probably caused by translocation phenomena. Based on the data obtained from the species within the studied genera, it is conceivable that variations in chromosomes, both structurally and in the distribution of constitutive heterochromatin, exert a significant influence on the evolution of the karyotype. Moreover, in many entities belonging to the Neottieae tribe, these processes may also contribute to the diversification of the phenotype in some instances.

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.

Summary: Stable isotope signatures of fungal sporocarps have been instrumental in identifying carbon gains of chlorophyllous orchids from a fungal source. Yet, not all mycorrhizal fungi produce macroscopic sporocarps and frequently fungi of different taxa occur in parallel in orchid roots.To overcome this obstacle, we investigated stable isotope signatures of fungal pelotons extracted from orchid roots and compared these data to the respective orchid and reference plant tissues. Anoectochilus sandvicensis and Epipactis palustris represented specialized or unspecialized rhizoctonia‐associated orchids. Epipactis atrorubens and Epipactis leptochila are orchids considered ectomycorrhiza‐associated with different preferences for Basidio‐ and Ascomycota.13C enrichment of rhizoctonia pelotons was minor compared with plant tissues and significantly lower than enrichments of pelotons from ectomycorrhizal Epipactis species. 15N values of pelotons from E. leptochila and E. atrorubens showed similar patterns as known for respective sporocarps of ectomycorrhizal Ascomycota and Basidiomycota, however, with an offset towards lower 15N enrichments and nitrogen concentrations.Our results suggest an explicit fungal nutrition source of orchids associated with ectomycorrhizal fungi, whereas the low 13C enrichment in rhizoctonia‐associated orchids and fungal pelotons hamper the detection of carbon gains from fungal partners. 15N isotopic pattern of orchids further suggests a selective transfer of 15N‐enriched protein‐nitrogen into orchids. [ABSTRACT FROM AUTHOR]

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Mountain forests of the Northern Tien Shan are distinguished by high floristic diversity and a large number of plant communities. Among the latter there are several unique relict plant communities dominated by Picea, Malus, Prunus, and Celtis spp, as well as tugai forests. The most species-rich are forests dominated by Picea schrenkiana Fisch. & C.A. Mey. (450-500 species), Malus sieversi (Ledeb.) M. Roem. (400 species), and Armeniaca vulgaris Lam. (about 250 species). The least species-rich are tugai forests with Hippophae rhamnoides L. (140 species) and hackberry forests dominated by Celtis caucasica Willd. (119 species). In these forests, there are 24 Red Book species, including Hepatica falconeri (Thoms.) Steward, Epipactis palustris (L.) Crantz, Tulipa tarda Stapf, and Gymnospermium altaicum (Pall.) Spach., which are the objects of special research. The article presents the results of the study of natural populations of the last three species, in particular, the numbers and density, age spectra and morphological variability of generative individuals. For one of the rarest species, Hepatica falconeri, the results of two years of observations of an introduced population established in the village of Saty near the central estate of Kolsay Koldery National Park. It was created by transferring generative individuals from the largest natural population (Taldy Gorge, 1693 m). Comparative data on the species phenology, including the relationship between the rate of flower opening, illumination and temperature, were obtained. The article emphasizes the lack of study of the distribution of certain rare species within the Northern Tien Shan, as evidenced by new findings of the authors: Epipactis palustris and Paeonia intermedia C.A. Mey. in the Kolsay Koldery National Park. Based on this, the authors propose to conduct special studies or include the study of rare species in the work plans of specially protected territories. [ABSTRACT FROM AUTHOR]

On the basis of numerous own observations, occasionally with information from third parties and with selected information from relevant literature, all orchid species of the island of Krk mentioned or recorded so far are described in detail. The basis for this is more than 2,000 entries of documented occurrences on the island of Krk, which were taken from our „data treasure“ with more than 12,000 entries on the orchid flora of the northeastern Adriatic (period of the years 1999 to 2022). The distribution maps, also based mainly on our own data, were supplemented by reports of findings in the Flora of Istria and, in some cases, with information from the Flora Croatica Database, thus providing an almost complete representation of the currently known distribution picture. [ABSTRACT FROM AUTHOR]

Abstract Background The flowers of some species of orchids produce nectar as a reward for pollination, the process of transferring pollen from flower to flower. Epipactis albensis is an obligatory autogamous species, does not require the presence of insects for pollination, nevertheless, it has not lost the ability to produce nectar, the chemical composition of which we examined by gas chromatography-mass spectrometry (GC–MS) method for identification of potential insect attractants. Results During five years of field research, we did not observe any true pollinating insects visiting the flowers of this species, only accidental insects as ants and aphids. As a result of our studies, we find that this self-pollinating orchid produces in nectar inter alia aliphatic saturated and unsaturated aldehydes such as nonanal (pelargonal) and 2-pentenal as well as aromatic ones (i.e., syringaldehyde, hyacinthin). The nectar is low in alkenes, which may explain the absence of pollinating insects. Moreover, vanillin and eugenol derivatives, well-known as important scent compounds were also identified, but the list of chemical compounds is much poorer compared with a closely related species, insect-pollinating E. helleborine. Conclusion Autogamy is a reproductive mechanism employed by many flowering plants, including the orchid genus Epipactis, as an adaptation to growing in habitats where pollinating insects are rarely observed due to the lack of nectar-producing plants they feed on. The production of numerous chemical attractants by self-pollinated E. albensis confirms the evolutionary secondary process, i.e., transition from ancestral insect-pollinating species to obligatory autogamous.

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Background: The flowers of some species of orchids produce nectar as a reward for pollination, the process of transferring pollen from flower to flower. Epipactis albensis is an obligatory autogamous species, does not require the presence of insects for pollination, nevertheless, it has not lost the ability to produce nectar, the chemical composition of which we examined by gas chromatography-mass spectrometry (GC-MS) method for identification of potential insect attractants. Results: During five years of field research, we did not observe any true pollinating insects visiting the flowers of this species, only accidental insects as ants and aphids. As a result of our studies, we find that this self-pollinating orchid produces in nectar inter alia aliphatic saturated and unsaturated aldehydes such as nonanal (pelargonal) and 2-pentenal as well as aromatic ones (i.e., syringaldehyde, hyacinthin). The nectar is low in alkenes, which may explain the absence of pollinating insects. Moreover, vanillin and eugenol derivatives, well-known as important scent compounds were also identified, but the list of chemical compounds is much poorer compared with a closely related species, insect-pollinating E. helleborine. Conclusion: Autogamy is a reproductive mechanism employed by many flowering plants, including the orchid genus Epipactis, as an adaptation to growing in habitats where pollinating insects are rarely observed due to the lack of nectar-producing plants they feed on. The production of numerous chemical attractants by self-pollinated E. albensis confirms the evolutionary secondary process, i.e., transition from ancestral insect-pollinating species to obligatory autogamous. [ABSTRACT FROM AUTHOR]

The orchid genera Epipactis Zinn and Ophrys L. are well-known by their complicated taxonomy and extensive debates over species richness within them. These genera are represented in Turkmenistan by two species each. Two of them, namely E. turcomanica K. P. Popov et Neshat. and O. kopetdagensis K. P. Popov et Neshat., were hitherto accepted as species endemic to Turkmenistan. In the present paper, these taxa are synonymized with broadly distributed E. persica (Soó) Hausskn. ex Nannf. and O. oestrifera M. Bieb., respectively. Thus, the genus Epipactis is represented in Turkmenistan by E. persica and E. veratrifolia Boiss. et Hohen., and the genus Ophrys is represented by O. mammosa Desf. and O. oestrifera. There are no species of orchids endemic to Turkmenistan.

While there is mounting evidence that ongoing changes in the climate system are shifting species ranges poleward and to higher altitudes, responses to climate change vary considerably between species. In general, it can be expected that species responses to climate change largely depend on how broad their ecological niches are, but evidence is still scant. In this study, we investigated the effects of predicted future climate change on the availability of suitable habitat for 14 Epipactis (Orchidaceae) species, and tested whether habitat specialists would experience greater changes in the extent of their habitats than habitat generalists. We used Maxent to model the ecological niche of each species in terms of climate, soil, elevation and land-use and projected it onto climate scenarios predicted for 2061–2080. To test the hypothesis that temperate terrestrial orchid species with small ranges or small niche breadths may be at greater risk under climate change than species with wide ranges or large niche breadths, we related niche breadth in both geographic and environmental space to changes in size and location of suitable habitat. The habitat distributions of half of the species shifted northwards in future projections. The area of suitable habitat increased for eight species but decreased for the remaining six species. If expansion at the leading edge of the distribution was not possible, the area of suitable habitat decreased for 12 species. Species with wide niche breadth in geographic space experienced greater northwards expansions and higher habitat suitability scores than species with small niche breadth. Niche breadth in environmental space was not significantly related to change in habitat distribution. Overall, these results indicate that terrestrial orchid species with a wide distribution will be more capable of shifting their distributions under climate change than species with a limited distribution, but only if they are fully able to expand into habitats at the leading edge of their distributions.

We report on observations of four native Epipactis species in the Lesse et Lomme region of southern Belgium. Their morphological and ecological traits are described and two hybrids between E. helleborine and E. purpurata on the one hand, E. muelleri, on the other hand, are discussed. Also presented is a singular plant, which we call “l’épipactis femelle » (the female helleborine), probably a hybrid between E. muelleri and E. neglecta, which is totally devoid of male reproductive parts. Finally, we briefly discuss the role of forests, coniferous as well as deciduous, in favouring the development and evolution of species such as the members of genus Epipactis that inhabit them. [ABSTRACT FROM AUTHOR]

Epipactis albensis Novâkovâ & Rydlo (Orchidaceae), a species new to the flora of Lithuania, was found in the southwestern part of the country, in the Bukta Forest (the Zuvintas Biosphere Reserve, Marijampole district) in 2020, and the nearby Dalgine Forest (the Dalgines Botanical Reserve) in 2021. The closest isolated population is located ca. 340 km southwest, and newly recorded populations form an island outside the main distribution area. Species populations occur primarily in the Tilio-Carpinetum betuli plant community showing ecological plasticity. A part of the population occurs in habitats with anthropogenic influence. Epipactis albensis is the only obligate autogamous Epipactis species known in Lithuania until now. Although its flowering season starts at the end of July or the beginning of August, solitary flowering specimens can be found in September or even later. According to the assessment applying the IUCN criteria, species should be treated as endangered (EN) [B2a,b(ii)c(iv) D] in Lithuania. [ABSTRACT FROM AUTHOR]

Using proteomics-based identification by matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS), we conducted the first analysis of the composition of endophytic bacteria isolated from different parts of selected Epipactis species, i.e. the buds, the inflorescences and the central part of the shoots, as well as the rhizomes. We identified aerobic and anaerobic bacteria, including such taxa as Bacillus spp., Clostridium spp., Pseudomonas spp. and Stenotrophomonas spp., which may be considered as promoting plant growth. Because most of the indicated bacteria genera belong to spore-producing taxa (spores allow bacterial symbionts to survive adverse conditions), we suggest that these bacteria species contribute to the adaptation of orchids to the environment. We found clear differences in the microbiome between investigated closely related taxa, i.e., Epipactis albensis, E. helleborine, E. purpurata and E. purpurata f. chlorophylla. Some of the analysed orchid species, i.e. E. albensis and E. purpurata co-occur in habitats, and their bacterial microbiomes differ from each other. [ABSTRACT FROM AUTHOR]

The static properties of leaves with parallel venation from terrestrial orchids of the genus Epipactis were modelled as coupled elastic rods using the geometrically exact Cosserat theory and the resulting boundary-value problem was solved numerically using a method from Shampine, Muir and Xu. The response of the leaf structure to the applied force was obtained from preliminary measurements. These measurements allowed the Young's modulus of the Epipactis leaves to be determined. The appearance of wrinkles and undulation characteristics for some leaves has been attributed to the small torsional stiffness of the leaf edges. [ABSTRACT FROM AUTHOR]

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Background and Aims The geographical distribution of plant species is linked fundamentally not only to environmental variables, but also to key traits that affect the dispersal, establishment and evolutionary potential of a species. One of the key plant traits that can be expected to affect standing genetic variation, speed of adaptation and the capacity to colonize and establish in new habitats, and therefore niche breadth and range size, is the plant mating system. However, the precise role of the mating system in shaping range size and niche breadth of plant species remains unclear, and different studies have provided contrasting results. In this study, we tested the hypothesis that range size and niche breadth differed with mating system in the orchid genus Epipactis. Methods We modelled the ecological niches of 14 Epipactis species in Europe using occurrence records and environmental satellite data in Maxent. Niche breadth and niche overlap in both geographic and environmental space were calculated from the resulting habitat suitability maps using ENMTools, and geographic range was estimated using α-hull range definition. Habitat suitability, environmental variable contributions and niche metrics were compared among species with different mating systems. Key Results We did not detect significant differences in niche breadth, occurrence probability or geographical range between autogamous and allogamous Epipactis species, although autogamous species demonstrated notably low variation in niche parameters. We also found no significant differences in niche overlap between species with the same mating system or different mating systems. For all Epipactis species, occurrence was strongly associated with land cover, particularly broad-leafed and coniferous forests, and with limestone bedrock. Conclusions These results suggest that the mating system does not necessarily contribute to niche breadth and differentiation, and that other factors (e.g. mycorrhizal specificity) may be more important drivers of range size and niche breadth in Epipactis and orchids in general. [ABSTRACT FROM AUTHOR]

Aim: Interactions with mycorrhizal fungi are increasingly recognized as an important factor underlying the distribution and abundance of orchid species. However, the geographical distribution of orchid mycorrhizal fungi (OMF) and how their communities vary over large geographical areas are less well understood. Because climatic and environmental similarity may decrease with geographical distance or because some OMF have limited dispersal capabilities, similarities in orchid mycorrhizal communities can be expected to decrease with increasing distances separating orchid populations. However, up till now empirical evidence is largely lacking. Location: Eurasia. Taxa: Gymnadenia conopsea (L.) R. Brown and Epipactis helleborine (L.) Crantz. Methods: High‐throughput sequencing was used to perform a cross‐continental comparison of OMF that associate with two widespread Eurasian terrestrial orchids, Epipactis helleborine and Gymnadenia conopsea. Both phylogenetic and nonphylogenetic measures of community dissimilarity and their components were calculated and related to geographical distances using Mantel tests. Results: Our results showed that in both orchid species similarity in mycorrhizal communities decreased significantly with geographical distance. Decomposing the contribution of spatial turnover and nestedness to overall dissimilarity showed that the observed dissimilarity was mainly the result of species replacement between regions, and not of species loss. Similarly, a strong relationship was observed between phylogenetic community dissimilarity and geographical distance. Decomposing PCD values into a nonphylogenetic and phylogenetic component showed that orchid populations located closely next to each other were likely to contain the same operational taxonomic units (OTUs), but that the non‐shared taxa came from different phylogenetic clades. Species indicator analyses showed that the majority of OMF OTUs were restricted to particular geographical areas. However, some OTUs occurred in both continents, indicating that some fungi have very wide distributions. Main conclusions: Overall, these results demonstrate that orchid mycorrhizal communities differ substantially across large geographical areas, but that the distribution of orchids is not necessarily restricted by the distribution of particular OMF. Hence, widespread orchid species can be considered mycorrhizal generalists that are flexible in the OMF with which they associate across large geographical areas. [ABSTRACT FROM AUTHOR]

Epipactis tallosii is considered as one of the highly threatened European orchid species due to its local distribution and small isolated populations that are characterized by decreasing trends. The species is now enlisted in the endangered (EN) category of the Red List. Nevertheless, during the last decade, multiple new populations of the species were found in Hungary, while our field surveys indicated that E. tallosii populations regularly occur in poplar (Populus) plantations. Here we conducted a thorough field survey of poplar plantations, by visiting 182 plantations in Hungary and surrounding countries in the Pannonian Biogeographic Region. We found E. tallosii in 23% of the visited plantations, and counted a total of c. 4000 generative shoots. Moreover, we documented the occurrence of the species to four countries, where it has not been reported before: Croatia, Romania, Serbia and Ukraine. Alien and indigenous/mixed poplar plantations were similarly likely to harbour populations of E. tallosii. Our study suggests that the distribution area and number of populations of E. tallosii is much larger than previously assumed, and that poplar plantations serve as suitable habitat islands in the agricultural landscapes for this orchid. In the light of our results, we suggest the reassessment of the IUCN category of E. tallosii and to re-categorize it as Near Threatened (NT). Citation: Süveges K., Löki V., Lovas-Kiss Á., Ljubka T., Fekete R., Takács A., Vincze O., Lukács B. A. & Molnár V. A. 2019: From European priority species to characteristic apophyte: Epipactis tallosii (Orchidaceae). – Willdenowia 49: 401–409. doi: https://doi.org/10.3372/wi.49.49310 Version of record first published online on 4 December 2019 ahead of inclusion in December 2019 issue. [ABSTRACT FROM AUTHOR]

The Epipactis helleborine (L.) Crantz group is one of the most taxonomically challenging species complexes within the genus Epipactis. Because of the exceptionally high levels of morphological variability and the ability to readily cross with other species, ninety different taxa at various taxonomic ranks have already been described within its nominative subspecies, but the taxonomic status of most of them is uncertain, widely disputed, and sometimes even irrelevant. The present review is based on results of the most recent research devoted to the E. helleborine group taxonomy. In addition, we analysed data about taxa belonging to this group presented in some research articles and monographs devoted directly to the genus Epipactis or to orchids in certain area(s). Based on the reviewed literature and data collected in four taxonomic databases available online, we propose an updated list of the 10 currently accepted taxa in the E. helleborine group (two species, six subspecies, and two varieties), which includes E. helleborine (L.) Crantz subsp. helleborine; E. helleborine subsp. bithynica (Robatsch) Kreutz; E. helleborine subsp. distans (Arv.-Touv.) R.Engel and P.Quentin; E. helleborine subsp. neerlandica (Verm.) Buttler; E. helleborine var. tangutica (Schltr.) S.C.Chen and G.H.Zhu; E. helleborine subsp. tremolsii (Pau) E.Klein; E. helleborine subsp. voethii (Robatsch) Jakubska-Busse, Żołubak, and Łobas, stat. nov.; E. condensata Boiss. ex D.P.Young; E. condensata var. kuenkeleana (Akhalk., H.Baumann, R.Lorenz, and Mosul.) Popovich; and E. cupaniana C.Brullo, D’Emerico, and Pulv.

The article informs that mixotrophic orchids such as Epipactis and Cephalanthera do not use photosynthesis for perennial underground organs. It mentions that orchids roots associate with ectomycorrhizal fungi that themselves associate with the surrounding trees, which are the ultimate carbon source of the whole consortium.

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The analysis of flowers collected at different stages of anthesis provides strong evidence to conclude that the shell-shaped hypochile and the knobs of epichile form a nectary. The scent comes from the aromatic constituents of nectar and the epichile tissue and the apices of all tepals (osmophores). The comparison between pollinated and unpollinated flowers revealed that the anthesis of unpollinated flowers lasted up to the 16th day. The nectariferous secretory cells formed single-layered epidermis and several layers of underlying parenchyma built by small, isodiametric cells with thin walls and dense cytoplasm, relatively large nuclei, supplied by collateral vascular bundles. During the floral lifespan, the residues of secreted material were higher on the hypochile cells. The lipoid-carbohydrate material and lipid globules in the cell walls and in the cytoplasm were localised. The abundance of starch grains was observed at the beginning of anthesis and their gradual reduction during the flower lifespan. At the end of anthesis in unpollinated flowers, the lipoid-carbohydrate-phenolic materials have been demonstrated. The phenolic material was the same as in plastoglobuli. The features such as irregular plasmalemma, the secretory vesicles that fuse with it, fully developed dictyosomes, numerous profiles of ER indicate vesicle-mediated process of secretion. The substances could be transported by vesicles to the periplasmic space via granulocrine secretion and then to the external surface. Both micro-channels and slightly developed periplasmic space were visible in the hypochile epidermis. This is the first time for anatomical survey of secretory tissue in pollinated and unpollinated flowers of E. helleborine. [ABSTRACT FROM AUTHOR]

Analysis of the seed morphology is a widely used approach in ecological and taxonomic studies. In this context, intraspecific variability with respect to seed morphology (size, weight, and density) was assessed in two close Epipactis tremolsii Pau. populations sharing the same ecological conditions, except for the soil pollution distinguishing one of them. Larger and heavier seeds were found in plants growing on the heavy metal polluted site, while no differences in seed density were detected between seeds produced by plants growing on the contaminated and the control site. Moreover, seed coats and embryos varying together in their dimensions were described in the control population, while coats varying in their size independently from embryos were described in plants growing on the polluted site. Seeds from the two studied populations significantly differed in several parameters suggesting that intraspecific seed variability occurred in the case study.

A new infraspecific taxon in Orchidaceae, Epipactis purpurata f. variegata Żołubak & Jakubska-Busse f. nova, with variegated leaves, is described and illustrated from Lower Silesia, Poland. [ABSTRACT FROM AUTHOR]

The rewarding orchid Epipactis flava was studied in NW Thailand. Its flowers were visited by a wide range of insects, most of which served as pollinators. The most frequent pollen bearers were (in decreasing order): the cricket Homoeoxipha lycoides, stingless bees of the Tetragonula testaceitarsis/hirashimai complex, hoverflies of subfam. Syrphinae, the wasp Polybioides gracilis and sweat bees of subfam. Halictinae. We found no evidence of a link between the rheophytic habit of E. flava and its pollinator fauna. Whereas most pollinators visited the flowers to feed on nectar, females of Episyrphus alternans (Syrphidae: Syrphinae) were observed to oviposit despite the absence of prey for their young. Hence, we suggest that dual pollination systems contribute to the opportunist strategy of E. flava, and we discuss, in a phylogenetic framework, how the strategy fits in with those previously reported for Epipactis sect. Arthrochilium. The elastic attachment of the epichile (a universal trait in sect. Arthrochilium) was found to promote outcrossing, and we hypothesize that loss of the elastic hinge has provided a key innovation facilitating recurrent evolution of obligate autogamy in sect. Epipactis (which is nested in sect. Arthrochilium). [ABSTRACT FROM AUTHOR]

In this contribution, the conservation status assessment of four vascular plants according to IUCN categories and criteria are presented. It includes the assessment of Epipactis maricae (Croce, Bongiorni, De Vivo & Fori) Presser & S.Hertel at global level, and the regional assessment of Cerinthe retorta Sm. (Italy), Platanthera kuenkelei H.Baumann subsp. kuenkelei (Europe) and Typha elephantina Roxb. (Egypt). [ABSTRACT FROM AUTHOR]

Epipactis veratrifolia is recorded for the first time from Jebel Hafeet, a mountain in Abu Dhabi, United Arab Emirates (UAE). This species had been previously recorded from the northern part of the UAE and other areas on the Arabian Peninsula. The occurrence of E. veratrifolia in Jebel Hafeet expands the eastern distribution limits of the species and underlines the importance of biodiversity in the region. The new record fills a gap in the distribution of this species on the Arabian Peninsula. [ABSTRACT FROM AUTHOR]

Four Epipactis species are recorded as new taxa for the Romanian Flora: Epipactis nordeniorum Robatsch, E. tallosii A.Molnár & Robatsch, E. exilis P.Delforge and E. pontica Taubenheim. With this contribution, not only the number of Epipactis species in Romania is increased to fourteen, but also we fill in gaps in geographical distribution of Epipactis species at European level. For each investigated taxon, we provide information about the habitat type, associated plant species, and population size. [ABSTRACT FROM AUTHOR]

Reproductive isolation caused by divergent natural selection arising from differences between ecological environments or ecological interactions represents a major mechanism contributing to speciation, but its relative importance is poorly known. In this study, controlled reciprocal crossings and seed germination experiments were combined with genetic and morphometric analyses to test the hypothesis that previously described differences in mycorrhizal communities between forest and dune ecotypes of Epipactis helleborine were sufficiently strong to create complete reproductive isolation between the two ecotypes. Molecular analyses using 770 SNP markers showed that the two ecotypes were genetically distinct and that populations of the dune ecotype were genetically impoverished compared to populations of the forest ecotype. Morphologically, the two ecotypes were also significantly different, with plants of the dune ecotype generally being smaller than plants of the forest ecotype. The results further showed that immigrant seeds had a significantly lower probability of protocorm formation than native seeds, indicating strong immigrant inviability. Although both ecotypes were able to cross easily and to produce a large number of viable seeds, hybrid seeds showed significantly lower protocorm formation than pure seeds, further contributing to reproductive isolation. Overall, these results indicate that interfertile populations of a widespread orchid adapting to contrasting environments diverge as a consequence of concurrent selection acting against immigrants and hybrids and suggest that mycorrhizal fungi can play a role in the early stages of plant speciation. [ABSTRACT FROM AUTHOR]

The selection and validation of proper distinguishing characters are of crucial importance in taxonomic revisions. The modern classifications of orchids utilize the molecular tools, but still the selection and identification of the material used in these studies is for the most part related to general species morphology. One of the vegetative characters quoted in orchid manuals is leaf arrangement. However, phyllotactic diversity and ontogenetic changeability have not been analysed in detail in reference to particular taxonomic groups. Therefore, we evaluated the usefulness of leaf arrangements in the taxonomy of the genus Epipactis Zinn, 1757. Typical leaf arrangements in shoots of this genus are described as distichous or spiral. However, in the course of field research and screening of herbarium materials, we indisputably disproved the presence of distichous phyllotaxis in the species Epipactis purpurata Sm. and confirmed the spiral Fibonacci pattern as the dominant leaf arrangement. In addition, detailed analyses revealed the presence of atypical decussate phyllotaxis in this species, as well as demonstrated the ontogenetic formation of pseudowhorls. These findings confirm ontogenetic variability and plasticity in E. purpurata. Our results are discussed in the context of their significance in delimitations of complex taxa within the genus Epipactis.

The group of Epipactis helleborine s.l. includes several subspecies. A new subspecies Epipactis helleborine subsp. moratoria was determined in Slovenia in 2015. It thrives in mixed wood in a region of Gorica at Raztez. Morphologic and phenomenological comparison confirmed clear differences between E. helleborine subsp. moratoria and E. helleborine subsp. helleborine. The characteristic differences seen in E. helleborine subsp. moratoria are the stem which is more or less bent at the level of the leaf base, the plants are smaller and more slender than E. helleborine, there are also differences in the flowers and the leaves. The ovary in E. moratoria is often in a horizontal position, especially at the time of fruiting, wheras in E. helleborine ovary usually hangs down. All of the wild orchids in Slovenia are protected species and among them it is Epipactis helleborine subsp. moratoria which, up till now is only known at one site. [ABSTRACT FROM AUTHOR]

A new natural hybrid Epipactis × subtilis Jakubska-Busse, Żołubak & Łobas, hybr. nat. nov. ( Epipactis albensis × Epipactis purpurata) is described and documented from the province of Lower Silesia (SW Poland). The hybrid's morphological characteristics are compared with those of the parental species. [ABSTRACT FROM AUTHOR]