C and N stable isotope signatures reveal constraints to nutritional modes in orchids from the Mediterranean and Macaronesia

We compared the nutritional modes and habitats of orchids (e.g., autotrophic, partially or fully mycoheterotrophic) of the Mediterranean region and adjacent islands of Macaronesia. We hypothesized that ecological factors (e.g., relative light availability, surrounding vegetation) determine the nutritional modes of orchids and thus impose restrictions upon orchid distribution. Covering habitats from dark forests to open sites, orchid samples of 35 species from 14 genera were collected from 20 locations in the Mediterranean and Macaronesia to test for mycoheterotrophy. Mycorrhizal fungi were identified via molecular analyses, and stable isotope analyses were applied to test whether organic nutrients are gained from the fungal associates. Our results show that orchids with partial or full mycoheterotrophy among the investigated species are found exclusively in Neottieae thriving in light-limited forests. Neottioid orchids are missing in Macaronesia, possibly because mycoheterotrophy is constrained by the lack of suitable ectomycorrhizal fungi. Furthermore, most adult orchids of open habitats in the Mediterranean and Macaronesia show weak or no N gains from fungi and no C gain through mycoheterotrophy. Instead isotope signatures of some of these species indicate net plant-to-fungus C transfer.     

Niche analysis of orchids of serpentine and non-serpentine areas: Implications for conservation

Orchids are known for their species richness, intriguing ecology, rarity and the fact that they grow in almost all terrestrial ecosystems. Although numerous studies about their ecology have been carried out concerning calcareous areas, little is known about orchids that occur in serpentine habitats. The aim of this study was to investigate the ecological preferences of orchids in serpentine and non-serpentine areas on the model of the Valjevo Mountain Range (W Serbia). Niche analysis of orchids was performed using outlying mean index analysis. Data concerning geographical coordinates, altitude, habitat type, inclination, bedrock type, light regime, soil moisture, acidity, nitrogen and temperature were used as explanatory variables. Data of 33 orchid taxa from 407 localities were analysed. The most important gradients that govern orchid distribution were geological bedrock, light regime and temperature. The results have shown that only Anacamptis morio and Gymnadenia conopsea have statistically significantly larger populations on serpentine compared with non-serpentine bedrocks. This study highlights the importance of serpentine habitats as orchid habitats, bearing in mind the occurrence of rare species and species which were found exclusively in serpentine habitats.     

西双版纳片段化石灰岩森林附生兰科植物多样性研究

西双版纳石灰岩地区拥有丰富的兰科植物资源,但近年来随着橡胶树的大面积种植,使得该地区很多石灰岩森林呈片段化。为了解片段化石灰岩森林中附生兰科植物多样性状况以及片段化对附生兰科植物的影响,该研究选取了生境片段化的青岩寨和曼纳览,以及连续生境的绿石林和巴卡新寨等4个样地的29个样方进行多样性调查和对比研究。结果表明:4个石灰岩森林样共记录到附生兰科植物34属76种1 528株(丛)。通过对坡向、坡度、海拔、地形、郁闭度和森林类型等6个环境因子与附生兰科植物丰富度进行CCA分析,发现石灰岩地区附生兰科植物的分布主要受海拔和森林类型2个因素的影响,在海拔较高的青岩寨和巴卡新寨附生兰科植物物种丰富度高于低海拔的绿石林和曼纳览,而片段化对附生兰科植物的物种多样性和多度均无显著影响,这可能与片段化的历史较短有关。虽然目前来看生境片段化对石灰岩地区的附生兰科植物多样性无显著影响,但生境脆弱的石灰岩森林植被的保护对于兰科植物多样性保护则更具重要性。     

Generalism in the interaction of Tulasnellaceae mycobionts with orchids characterizes a biodiversity hotspot in the tropical Andes of Southern Ecuador

Biotic interactions play an important role in the assembly and stability of communities. All orchids depend on mycobionts for early establishment, but whether individual orchid species depend on a specific or broad spectrum of mycobionts is still a matter of debate. Tulasnellaceae (Basidiomycota) is the richest and most widespread mycobiont worldwide. We assessed Tulasnellaceae richness in epiphytic and terrestrial orchids in different habitats, and evaluated the degree of generalism in orchid-Tulasnellaceae interactions and the robustness of this mutualistic system to the extinction of mycobiont partners. We sampled 114 orchid individuals including all common and rare species in 56 plots of 1 m² in 3 habitats: pristine forest, regenerating forest and a landslide site in a tropical montane rainforest in Southern Ecuador. We found 52 orchid and 29 Tulasnellaceae species. The composition of Tulasnellaceae OTUs was moderately to highly similar across habitats and between orchid growth forms. A significantly nested network architecture indicated the existence of a core of generalist Tulasnellaceae OTUs interacting with both rare and common orchids. Terrestrial and epiphytic orchids showed significant differences in robustness to the extinction of their Tulasnellaceae mycobionts. Thus, generalist mycobionts may be relevant for the preservation of hyperdiverse orchid communities in the tropics.     

Vegetation factors influencing the establishment and growth of the endangered Japanese orchid,Cypripedium macranthos var. rebunense

Cypripedium macranthos var. rebunense is a well-known endangered terrestrial orchid that is endemic to the cool-temperate Rebun Island off the coast of northwestern Hokkaido, Japan. The present study aimed to identify suitable methods for conservation of this orchid through management of the natural vegetation. We analyzed the vegetation types and plants growing sympatrically with this orchid using two-way indicator species analysis (TWINSPAN) and detrended correspondence analysis (DCA) ordination. Reproductive orchids can survive in various broad-leaved and short to medium-height herbaceous meadows with high plant species richness at a small scale in coastal regions. In order to maintain C. macranthos var. rebunense populations, succession to dense and tall large-leaved herbs, Sasa scrub, shrubs, and trees, should be suppressed. Favorable vegetation for the establishment of seedlings of this orchid includes narrow-leaved, medium-sized grasses, sedges, herbs, mosses, and prostrate mat-forming shrubs. The ground cover offered by these plants and their litter may favor stability of moisture, temperature, and suitable light conditions at the ground surface. In situ seed sowing at such ‘safe sites’ is a potential low-impact technique to enhance seedling recruitment of this orchid.     

Epiphytism and pollinator specialization: drivers for orchid diversity?

Epiphytes are a characteristic component of tropical rainforests. Out of the 25,000 orchid species currently known to science, more than 70% live in tree canopies. Understanding when and how these orchids diversified is vital to understanding the history of epiphytic biomes. We investigated whether orchids managed to radiate so explosively owing to their predominantly epiphytic habit and/or their specialized pollinator systems by testing these hypotheses from a statistical and phylogenetic standpoint. For the first approach, species numbers of 100 randomly chosen epiphytic and terrestrial genera were compared. Furthermore, the mean number of pollinators per orchid species within the five subfamilies was calculated and correlated with their time of diversification and species richness. In the second approach, molecular epiphytic orchid phylogenies were screened for clades with specific suites of epiphytic adaptations. Epiphytic genera were found to be significantly richer in species than terrestrial genera both for orchids and non-orchids. No evidence was found for a positive association between pollinator specialization and orchid species richness. Repeated associations between a small body size, short life cycle and specialized clinging roots of twig epiphytes in Bulbophyllinae and Oncidiinae were discovered. The development of twig epiphytism in the first group seems repeatedly correlated with speciation bursts.     

三种杓兰根相关真菌多样性和生态功能

【背景】除了菌根真菌(Orchid mycorrhizal fungi,OrMF)外,兰科植物根中还有其它内生真菌,称为根相关真菌(Root-associated fungi,RAF)。【目的】采用分离培养的方法获得同一栖息地针叶林和灌木林两种不同生境西藏杓兰、黄花杓兰和无苞杓兰的RAF菌株,研究其真菌谱系、多样性和生态功能结构。【方法】从杓兰根碎屑中分离RAF,通过总DNA提取、PCR扩增及测序得到ITS(Internaltranscribedspacer)序列;进行系统发育和多样性分析,并通过NCBI数据库比对得到相似性最高序列的注释信息来分析RAF生态学特性。【结果】共分离得到278株RAF,25种OTU类型,包括23个子囊菌门OTU,2个毛霉菌门OTU。RAF物种丰富度分析发现西藏杓兰的较黄花杓兰高,不同生境没有显著差异;不同杓兰物种较不同生境的RAF群落分化程度高。生态功能分析显示25个OTU包括共生型、腐生型和致病型3种营养型,以及外生菌根菌群、植物病原菌群、内生真菌群、动物病原菌群、真菌寄生菌群、杜鹃花类菌根群、未定义的腐生菌群和不确定型8种共位群。【结论】阐明不同生境采集的不同杓兰中RAF的分布特点和生态功能,为未来研究RAF与杓兰属植物的共生关系奠定基础。     

Is orchid species richness and abundance related to the conservation status of oak forest?

The response of organisms to anthropogenic or natural modification of the environment is one of the most important questions in conservation biology and ecological theory. In spite of the fact that orchids are one of the most studied groups of plants, little information exists regarding their response to habitat alteration. The few existing studies are biased toward European orchid species and no consensus exists with regard to the response of orchids to human and natural disturbance. In this study, we sampled 30 transects (0.1 ha each) of oak forest located in Morelos, Central Mexico, and measured 13 variables related to forest aging and stump abundance, and six variables of orchid species richness and abundance. Neither abundance nor the richness and specific abundance of orchid genus or species were related to timber extraction. The abundance of dead standing trees in the forest, a surrogate variable of forest age, was positively related to abundance of orchids of the genus Malaxis, orchid richness and orchid abundance. This finding suggests that the conservation of all facets of the studied forest orchid community is dependent on natural processes (such as self-thinning) and the maintenance of older areas of the forest, and concurs with previous studies that suggest that natural disturbance is a key process for orchid survival.     

Diversity, flower visitation frequency and generalism of pollinators in temperate rain forests of Chiloé Island, Chile

Species richness and taxonomic composition of pollinator assemblages are documented for 26 plant species from temperate rain forests of northern Chiloé Island, southern Chile (42°30'S). We investigated the patterns of generalism and specialization among plants and animal pollinators by comparing the flower visit frequency by different pollen vectors during the spring and summer months of three consecutive years (2000–2002). Species studied exhibited a range of floral morphologies (radial vs. zygomorphic, open vs. tubular) and rewards (nectar and/or pollen). Overall, we recorded 172 pollinator species, with an average of 6.6 species of pollen vectors/plant species. Pollinators visited an average of 15.2 plant species/pollen vector. Pollinator assemblages were dominated by Coleoptera (75 species), Diptera (56 species) and Hymenoptera (30 species), but passerine birds and hummingbirds were also important. The most specialized plants were vines, including the bee-pollinated genus Luzuriaga (Philesiaceae) and two endemic species of hummingbird-pollinated Gesneriaceae. Hymenoptera contributed 41.2% of all visits, with the bumblebee Bombus dalhbomii accounting for 22.5% of these. Plants with unspecialized flower morphology supported a higher species richness of pollinators, but visiting rates did not differ from specialized flowers.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 399–416.     

Poromitra crassiceps (Teleostei, Melamphaidae) associated with the 500 fathoms fauna off Argentina

We established the presence of Poromitra crassiceps on the slope off Argentina (49°29.8'S; 55°27.4'W, 941 m depth) during a research cruise in 1978. The report is based on a single specimen 130.09 mm total length.     

Untangling above‐ and belowground mycorrhizal fungal networks in tropical orchids

Orchids typically depend on fungi for establishment from seeds, forming mycorrhizal associations with basidiomycete fungal partners in the polyphyletic group rhizoctonia from early stages of germination, sometimes with very high specificity. This has raised important questions about the roles of plant and fungal phylogenetics, and their habitat preferences, in controlling which fungi associate with which plants. In this issue of Molecular Ecology, Martos et al. (2012) report the largest network analysis to date for orchids and their mycorrhizal fungi, sampling a total of over 450 plants from nearly half the 150 tropical orchid species on Reunion Island, encompassing its main terrestrial and epiphytic orchid genera. The authors found a total of 95 operational taxonomic units of mycorrhizal fungi and investigated the architecture and nestedness of their bipartite networks with 73 orchid species. The most striking finding was a major ecological barrier between above‐ and belowground mycorrhizal fungal networks, despite both epiphytic and terrestrial orchids often associating with closely related taxa across all three major lineages of rhizoctonia fungi. The fungal partnerships of the epiphytes and terrestrial species involved a diversity of fungal taxa in a modular network architecture, with only about one in ten mycorrhizal fungi partnering orchids in both groups. In contrast, plant and fungal phylogenetics had weak or no effects on the network. This highlights the power of recently developed ecological network analyses to give new insights into controls on plant–fungal symbioses and raises exciting new hypotheses about the differences in properties and functioning of mycorrhiza in epiphytic and terrestrial orchids.     

The Southern African orchid flora: composition, sources and endemism

Aim The Southern African orchid flora is taxonomically well known, but the biogeographical and diversity patterns have not yet been analysed. In particular, we want to establish whether (a) it is, like the Southern African flora in general, more diverse than would be expected from its latitude and area; (b) it is an African flora, or whether it contains palaeoendemic relicts of a Gondwanan orchid flora; (c) the diversity and endemism in the orchid flora is concentrated in particular biomes and habitat types; and (d) the patterns of endemism in the flora can be accounted for by current environmental parameters, or whether we need to invoke historical explanations. Location Southern Africa. Methods We used the recent floristic account of the Southern African orchids, in conjunction with a data base of over 14,642 herbarium records, to assign the species and subspecies of Southern African orchids to biomes, habitats, and clades. We explored the relationship between the number and endemism of entities (species, subspecies and varieties) and the biomes and habitats. We compared the richness of this flora with that of 31 other regions from all continents and latitudes, to establish whether the Southern African orchid flora is richer or poorer than expected. We assigned the Southern African orchid species to 16 monophyletic clades and mapped the global distribution of these clades to establish the continental affinities of the flora. Main conclusions The Southern African orchid flora is not any more diverse than could be expected from its latitude or area, while the two tropical African floras included were less diverse than expected. Latitude is an excellent predictor of regional orchid species richness; this might indicate that available habitat is more important for orchid diversity than gross area available, since latitude is probably correlated with the extent of suitable habitat. The Southern African orchid flora is clearly an African flora, since all clades are also found in tropical Africa, while many of them are absent from the Americas or Asia. Conversely, while most African orchid clades are also found in Southern Africa, both the Americas and Asia contain many clades absent from Africa. The distribution of orchid entities among the biomes in Southern Africa is very uneven, with two of the seven biomes totally devoid of orchids. Habitats and biomes that have no equivalent in tropical Africa are high in endemism, and habitats and biomes which are also well developed in tropical Africa are low in endemism. Endemism appears largely explained in terms of modern habitats. However, two patterns (the high endemism in the Succulent Karoo and the lack of endemism in the southern Cape among epiphytic orchids) may also be explained in terms of Quaternary climatic changes.     

Analysis of macrofungal communities reveals a complex reciprocal influence between Mediterranean montane calcareous grassland and surrounding forest habitats

Fungi are essential components of all terrestrial ecosystems. Despite the crucial ecological role of soil fungi in grasslands, knowledge about fungal community diversity and structure in Mediterranean meadow habitats is still fragmentary. We analyzed macrofungal communities in three geographically distinct Mediterranean montane calcareous grasslands and surrounding forests, by means of fruit body surveys. We investigated a number of biotic and abiotic factors influencing the studied fungal communities, including plant species composition. Out of 6365 fruit bodies, a total of 268 species belonging to 84 genera were found. In general, there was a significant correlation between plant species richness and fungal richness. Variation in vegetation and plant community structure accounted for approximately 20% of variance in fungal community structure. Tree and shrub vegetation played a dominant role in shaping the analyzed fungal communities, both in meadows and surrounding forests, with particular influence on ectomycorrhizal, litter, and lignicolous saprotrophic fungi. Fungal biodiversity in the studied meadows was increased by the presence of tree and shrub species from the adjacent forests, but was reduced by the increasing vegetation cover.     

广东象头山国家级自然保护区兰科植物垂直分布格局

于2019～2022年对广东象头山国家级自然保护区内的兰科植物进行专项调查,记录兰科植物种类、数量、生活型、海拔、生境等信息,分析保护区内兰科植物的垂直分布格局,并根据《广东高等植物红色名录》和《国家重点保护野生植物名录》对各海拔高度兰花濒危等级的物种数量进行统计。结果显示：(1)实地记录兰科植物33属48种,多数种类种群数量较少;(2)随着海拔上升,各海拔梯度的兰科植物总种数和不同生活型的种数均呈现“中间高度膨胀型”分布特点,峰值在中低海拔区域;(3)濒危物种主要集中在海拔200～500 m区间;(4)相邻海拔段的兰科植物种类相似性系数较高,垂直梯度差异越大,相似性系数越小。低海拔地区可能由于人为干扰频繁,兰科植物物种多样性较低;中海拔物种多样性最丰富;高海拔地区生长环境较苛刻,物种数量较少。     

Roadside verges and cemeteries: Comparative analysis of anthropogenic orchid habitats in the Eastern Mediterranean

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.     

Influence of mycorrhizal fungi on seed germination and growth in terrestrial and epiphytic orchids

Epiphytes constitute over 70% of orchid diversity, but little is known about the functioning of their mycorrhizal associations. Terrestrial orchid seeds germinate symbiotically in soil and leaf litter, whereas epiphytic orchids may be exposed to relatively high light levels from an early stage of development and often produce green seeds. This suggests that seedlings of the two groups of orchids may differ in their responses to light and requirements for mycorrhiza-supplied carbon. The interactive effects of light, exogenous carbon and mycorrhizal status on germination and growth were investigated in vitro using axenic agar microcosms for one tropical epiphyte and three geophytic orchid species. The geophytic species strongly depended on their mycorrhiza for growth and this could not be substituted by exogenous sucrose, whereas the epiphytic species achieved 95% of the mycorrhizal seedling volume when supplied with exogenous sucrose in the dark. Mycorrhiza status strongly interacted with light exposure, enabling germination. Light inhibited or severely reduced growth, especially for the terrestrial orchids in the absence of mycorrhiza. For the first time, this study showed the parallel ecological importance of mycorrhizal fungi in overcoming light inhibition of seed germination and growth in both terrestrial and epiphytic orchids.     

Composition, geographical affinities and endemism of the Iberian Peninsula orchid flora

The orchid flora of the Iberian Peninsula is relatively well known, but its biogeographical and diversity patterns have until now remained unanalysed. This work compares the richness of this flora with that of 27 other territories in different continents and at different latitudes, with the aim of establishing whether it is richer or poorer than might be expected. Latitude was found to be an excellent predictor of regional orchid species richness. With 122 taxa, the orchid flora of the Iberian Peninsula is more or less as diverse as that of other Mediterranean areas of similar latitude (e.g. France, Greece or Italy), but more diverse than other European or indeed North African orchid floras. In this study, the Iberian orchid species were assigned to eight monophyletic clades and the global distribution of these are mapped to establish continental affinities between the floras. A recent floristic account on the Iberian orchids was also used to assign the orchid taxa to habitats, and the relationship between the number of endemisms and their habitats was analysed. The patterns of endemism differed in different habitats. Very high levels of endemism were found in habitats peculiar to the Mediterranean Basin, indicating the relict status of its orchid flora.     

Studies of Mycorrhizal Fungi of Chinese Orchids and Their Role in Orchid Conservation in China—A Review

China has over 1,200 species of native orchids in nearly 173 genera. About one fourth of native species are of horticultural merit. Some species are of Chinese medicinal value. In fact, the demand on orchid species with high Chinese medicinal values such as Gastrodia elata, Dendrobium offcinale, along with demands on species of cultural importance, such as those in the genus of Cymbidium, is a major factor causing wild populations to diminish and in some cases, drive wild populations to the brink of extinction. These market demands have also driven studies on the role of mycorrhizal fungi in orchid seed germination, seedling and adult growth, and reproduction. Most of these mycorrhizal studies of Chinese orchids, however, are published in Chinese, some in medical journals, and thus overlooked by the mainstream orchid mycorrhizal publications. Yet some of these studies contained interesting discoveries on the nature of the mycorrhizal relationships between orchids and fungi. We present a review of some of these neglected publications. The most important discovery comes from the mycorrhizal studies on G. elata, in which the researchers concluded that those fungi species required to stimulate seed germination are different from those that facilitate the growth of G. elata beyond seedling stages. In addition, presence of the mycorrhizal fungi associated with vegetative growth of post-seedling G. elata hindered the germination of seeds. These phenomena were unreported prior to these studies. Furthermore, orchid mycorrhizal studies in China differ from the mainstream orchid studies in that many epiphytic species (in the genus of Dendrobium, as medicinal herbs) were investigated as well as terrestrial orchids (mostly in the genus Cymbidium, as traditional horticultural species). The different responses between epiphytic and terrestrial orchid seeds to fungi derived from roots suggest that epiphytic orchids may have a more general mycorrhizal relationship with fungi than do terrestrial orchid species during the seed germination stage. To date, orchid mycorrhizal research in China has had a strongly commercial purpose. We suggest that this continuing research on orchid mycorrhizal relationships are a solid foundation for further research that includes more rare and endangered taxa, and more in-situ studies to assist conservation and restoration of the endangered orchids. Knowledge on the identities and roles of mycorrhizal fungi of orchids holds one of the keys to successful restoration and sustainable use of Chinese orchids.     

Effects of altitude and climate in determining elevational plant species richness patterns: A case study from Los Tuxtlas,Mexico

Altitudinal changes of composition and richness of montane plant assemblages are complex, depending on the taxonomic group and gradient conditions, with different factors involved that are directly altitude-dependent (e.g., temperatures, air pressure) and altitude-independent (e.g., precipitation, cloud cover, area). In order to assess the relative impacts of temperature, precipitation, air humidity, and area of altitudinal belts on plant diversity, we analyzed diversity patterns of five species-rich groups, mostly herbaceous plants, in 74 forest plots along three climatically contrasting elevational transects from humid tropical lowland vegetation up to cloud forests at Los Tuxtlas, Mexico. We recorded 278 plant species, with ferns being the most species-rich group followed by orchids, bromeliads, aroids, and piperoids. The most striking results were the contrasting patterns and model results for terrestrial and epiphytic taxa. Whereas the richness of all terrestrial species taken together did not change significantly with elevation, vascular epiphytes showed increasing species numbers with altitude. However, a number of individual terrestrial taxa showed also significant elevation-related changes: aroids showed a marked decline with hight, orchids and piperoids increased, and ferns displayed a hump-shaped pattern with highest richness in mid-altitudes. Among the epiphytes, aroids declined while most other groups increased with altitude. This distinction is relevant for projections of responses of plant communities to climate change, which will lead to increased temperatures and to changing precipitation and cloud condensation regimes and thus will likely affect terrestrial and epiphytic species in different ways.