Diversity of mycorrhizal fungi of terrestrial orchids: compatibility webs, brief encounters, lasting relationships and alien invasions

The diversity of mycorrhizal fungi associated with an introduced weed-like South African orchid (Disa bracteata) and a disturbance-intolerant, widespread, native West Australian orchid (Pyrorchis nigricans) were compared by molecular identification of the fungi isolated from single pelotons. Molecular identification revealed both orchids were associated with fungi from diverse groups in the Rhizoctonia complex with worldwide distribution. Symbiotic germination assays confirmed the majority of fungi isolated from pelotons were mycorrhizal and a factorial experiment uncovered complex webs of compatibility between six terrestrial orchids and 12 fungi from Australia and South Africa. Two weed-like (disturbance-tolerant rapidly spreading) orchids — D. bracteata and the indigenous Australian Microtis media, had the broadest webs of mycorrhizal fungi. In contrast, other native orchids had relatively small webs of fungi (Diuris magnifica and Thelymitra crinita), or germinated exclusively with their own fungus (Caladenia falcata and Pterostylis sanguinea). Orchids, such as D. bracteata and M. media, which form relationships with diverse webs of fungi, had apparent specificity that decreased with time, as some fungi had brief encounters with orchids that supported protocorm formation but not subsequent seedling growth. The interactions between orchid mycorrhizal fungi and their hosts are discussed.     

Surprisingly high orchid diversity in travertine and forest areas in the Huanglong valley, China, and implications for conservation

The presence of such a large number of terrestrial orchid species in a small area (ca. 1 km²) of the Huanglong valley in southwestern China is uncommon for this country. Studying the relationship between the distribution patterns of these orchid species and their microenvironments may help us understand this uncommon phenomenon. We established 662 1 m × 1 m plots, measured the cover of each species and found that there were 33 orchid species distributed mainly in two different habitats, i.e. travertine areas and forest. In the travertine areas, 30 orchid species were found; the six most common ones being Cypripedium bardolphianum, Cypripedium flavum, Cypripedium tibeticum, Galearis diantha, Ponerorchis chusua and Phaius delavayi. However in the forested habitat, we found 21 orchid species; the most common ones being Tipularia szechuanica and Goodyera repens. Travertine areas had a higher number of orchid species as well as higher numbers of orchid species per plot as compared to forest. Light availability seems critical to the performance and distribution of orchid species. Stream flow through the travertine area during the orchids growing season appears to be an important factor in shaping and maintaining stable microenvironments favorable to the growth and reproduction of orchids. The results presented in this study suggest that some orchid species in the travertine area might be threatened if the travertine stream flows were to change or be disrupted.     

Long-term ecology of euglossine orchid-bees (Apidae: Euglossini) in Panama

Summary Abundance patterns during 6–7 years and orchid visitation were determined for 51 species of the 57 local euglossine bees. Male bees were counted at 3 chemical attractants presented in the same manner each month. Sites were separated by 75 km but included wet Atlantic forest at 500 m elevation, moist forest at 180 m near Barro Colorado Island, and cloud forest at 900 m near the Pacific ocean. 1. From 15 to 30 euglossine species of 4 genera were active in each month and site; monthly species number and general bee abundance were positively correlated. Many species had 3 annual abundance peaks (range 1–4) and were active throughout the year, but peak annual abundances rarely occurred during late wet or early dry seasons. In contrast, Eufriesea generally were present as adults only 1–2 months in a year. 2. Euglossine populations were exceptionally stable. Species at each site were more stable than any known insect population, and stability and abundance were positively associated. However, year-to-year population stability and the degree of seasonality were not correlated. Among the three sites, the more diverse (species rich) bee assemblages displayed lower stability; these were the wetter and upland sites. 3. The most abundant bees visited more orchid species. Eg. and El. each visited and average of 4 orchid species (range 0–13); Ex. and Ef. visited 0–3. Stable populations did not visit more or fewer orchid species than did unstable populations. 4. Less than 68% of species at each site visited orchid flowers; less than a few dozen of the 100–800 bees counted in a day carried orchid pollinaria. Over 20% of the euglossine species never were seen with pollinaria at any site and probably seldom visit orchids in central Panama. 5. Most bee species visited 1 or no fragrance orchids in a given habitat. Orchids tended to utilize common pollinators that seldom included more than 1 species, and they utilized stable or unstable, seasonal or aseasonal bees. However, the most stable and abundant bee, Eg. imperialis, rarely pollinated orchids; fewer than 10 of ca. 20000 bees carried pollinaria. 6. Orchids may interact primarily with discrete seasonal bee population peaks-probably the emerging adults. Although specialized orchid preferences are implicated for species that visit few or no local orchids but pollinate other species and carry pollinaria in other areas, euglossine bees do not need orchids to survive or reproduce.     

4. The role of plant growth regulators in the germination of forest tree seeds

To establish a role for PGRs in the germination of forest tree seeds, it must be demonstrated that PGRs are involved in the mechanisms that control the induction of and release from dormancy, and also that these mechanisms are operative in the seeds of woody plants. Four currently known concepts of seed dormancy are reviewed, citing the published evidence relating PGRs to tree seed germination. The germination of tree seeds appears to be controlled by a variety of external and internal factors. PGRs figure prominently among these factors, but the range of mechanisms by which such control is mediated may vary considerably—from physical to metabolic. Although significant support can be found for PGR involvement in the regulation of angiosperm seed dormancy, the evidence for gymnosperms is still inconclusive. This is primarily because our understanding of the control of germination—by PGRs or other means—is generally unsatisfactory. However, recent advancements in PGR technology, together with a more holistic concept of PGR action in biological systems, signal a more favourable climate and a revitalized approach to the investigation of the role of PGRs in plant growth and development.     

Fruit set, nectar reward, and rarity in the Orchidaceae

A review of comparative levels of reproductive success among nectariferous and nectarless orchids worldwide was compiled from a comprehensive survey of fruit set from 117 orchid species in the literature and from our own field studies. It confirms the hypothesis that nectariferous orchids are more successful in setting fruit than are nectarless species. Overall fruit set figures for nectarless and nectariferous orchids were 19.5 and 49.3% for North America, 27.7 and 63.1% for Europe, 41.4 and 74.4% for the temperate southern hemisphere, and 11.5 and 24.9% for the tropics, demonstrating that the dichotomy is consistent across all geographical areas. On average, the provision of nectar doubles the probability of fruit set in both temperate and tropical areas, but tropical orchids are remarkable in that all (whether nectarless or nectariferous, or terrestrial or epiphytic) display low fruit productivity (<50%). Fruiting failure in the tropics may be balanced by higher productivity per capsule, since tropical orchid fruits contain on average 150 times more seeds than temperate ones. Hybridization occurs more frequently among nectarless orchids in Britain and Europe than among nectariferous ones, and there is a significant positive association between orchid rarity and lack of nectar reward in the British Isles. Sexual reproduction in the Orchidaceae is predominantly pollinator dependent, but this can sometimes be successfully circumvented by asexual seed production (agamospermy) or, more frequently, by automatic self-pollination (autogamy). The proportion of highly successful nectarless orchids from all geographic areas is very low and comparable with that of orchids offering rewards other than nectar (～14% of species in each case) emphasizing that high reproductive success is only associated with nectar reward (53% of species). It is suggested that the evolution of nectar production within the family has been the most frequent means of escaping the reproductive limitations of low pollinator visitation frequencies.     

A Habitat Suitability Index (HSI) for the Western Prairie Fringed Orchid (Platanthera praeclara) on the Sheyenne National Grassland,North Dakota,USA

The Western Prairie Fringed Orchid (Platanthera praeclara) is a threatened species found on the Sheyenne National Grassland (SNG) in southeast North Dakota, USA. The SNG is subject to management for multiple uses including biodiversity conservation, livestock grazing and recreation. Therefore, there is a need for the development of indicators of suitable orchid habitat. The orchids are continuously monitored, but understanding of the relationship between landscape properties and orchid locations is limited. In this study data that characterize topography, moisture, and groundwater were used to construct indicators of landscape suitability and an overall Habitat Suitability Index (HSI) for the orchid. A LiDAR-derived DEM and groundwater well observations were used to develop landscape indicators. The Topographic Wetness Index (TWI: a measure of moisture on the landscape), the Topographic Position Index (TPI: a measure of position on the landscape), and the distance to groundwater (DTG: a measure of the distance from the land surface to the groundwater surface) provided the best set of indicators of orchid habitat. Point-based field observations of orchid occurrence were used to develop Orchid Suitability Metrics (OSMs) that identified the range of indicator values most strongly associated with orchids. These OSMs were used to define year by year suitability zones for each indicator that were combined to create the HSI. Comparison of orchid locations with groundwater elevations showed that orchids occurred on average 0.98 ± 0.39 (2σ) m above the water table. TWI and TPI demonstrated that orchids occur near flow paths and areas of lower elevation than their surroundings. HSI values of 0.67 and above were associated with 89.8% of all orchid observations used in the analysis. The landscape indicators, OSM concept and HSI could be generally applied to monitoring and conservation management of orchid habitat and the concept may be applicable to other valued species with similar niche properties.     

Infestation of the orchid Cephalanthera spp. by Parallelomma vittatum (Meigen, 1826) (Diptera: Scathophagidae) in Japan

It is important to understand the interactions between orchids and their herbivores, especially considering the rarity of most orchids and the potential negative effects of pests on orchid reproduction. Here, we report the infestation of leaves of Cephalanthera spp. orchids by Parallelomma vittatum (Scathophagidae) in Japan. The present study represents the first report of P. vittatum on orchid hosts in Asia.     

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.     

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.     

Wild orchid tuber collection in Iran: a wake-up call for conservation

Wild orchids are traditionally harvested as Salep and used in traditional medicine and ice-cream production in Iran. Recently however, illegal harvest of wild orchids for export appears to have grown. This study aimed to: (1) determine the diversity of harvested wild orchid species and their collection sites in Iran; and (2) study the current harvest status and trade chain and volume to estimate the total orchid plant extraction from natural populations. Field surveys of collectors and market surveys of traders were conducted to establish the diversity of collected species, to identify harvest hotspots, and to document harvesting and trade volumes. Sixteen species and subspecies from 7 genera of Orchidaceae are collected for their tubers. Based on estimates from the 2013 April to June harvest season more than 24.5 tons of fresh tubers were collected from three districts in Golestan province alone. It is estimated that this amount of tuber requires the lethal destructive harvesting of 5.5 –6.1 million orchids, with a market value of 320,000 USD. In the Tehran Bazar Salep trade during May–July 2013 was 1.9 tons of dried tubers, with estimated retail value of 310,000 USD. Current orchid collection practices in Iran, which have soared in recent years due to international demand, do not seem sustainable as all tubers are collected destructively. To preserve orchid populations, in the longterm, establishment of specific Orchid Conservation Areas and introduction of sustainable production practices, could alleviate harvesting pressure. In the midterm, development of a DNA barcoding-based molecular identification system could help to monitor and control illegal trade. In the near term, effective implementation of collection bans in excessively harvested areas and strengthening of current regulations are necessary to avoid the catastrophic effects of harvesting on orchid populations, as has been observed in Turkey.     

四川黄龙沟草本层植物种类组成与数量特征

四川黄龙沟草本植物群落在所调查样方中(N=662)共出现维管植物124种,分属于37科91属,有54个中国特有种:其中兰科植物多达21属33种,中国特有种有12个,新种一个。如此众多的地生兰科植物聚集在面积不到1公里的沟内,而且部分兰科植物种类在沟内形成优势草本群落,这在中国地生兰的分布区域中是不多见的。大多数物种,包括兰科植物在内,出现的频率都较小,并且分布不均匀。黄龙沟兰科植物主要分布在两种生境中,即钙化滩流地和森林中。在这两种生境中微环境条件以及兰科植物的种类组成、数量特征和分布格局的差异都很大。钙化滩流地的兰科植物种类数目和每个样方中包含的兰科植物种类数目都比森林生境中的高。钙化滩流地中发现有30种兰科植物,最为常见的是无苞杓兰、黄花杓兰、西藏杓兰、广布小碟兰、二叶根茎兰和少花鹤顶兰。森林生境中有21种兰科植物分布,其中筒距兰和斑叶兰出现最多,光照强度可能对兰科植物的分布起到重要的决定作用。钙化滩流地中的溪流对建立和维持兰科植物生存所必须的稳定环境条件起到至关重要的作用,如果随意改变水流的方向或减少水流的流量,将给某些兰科植物带来灾难性的后果。     

Atractiellomycetes belonging to the ‘rust’ lineage (Pucciniomycotina) form mycorrhizae with terrestrial and epiphytic neotropical orchids

Distinctive groups of fungi are involved in the diverse mycorrhizal associations of land plants. All previously known mycorrhiza-forming Basidiomycota associated with trees, ericads, liverworts or orchids are hosted in Agaricomycetes, Agaricomycotina. Here we demonstrate for the first time that Atractiellomycetes, members of the ‘rust’ lineage (Pucciniomycotina), are mycobionts of orchids. The mycobionts of 103 terrestrial and epiphytic orchid individuals, sampled in the tropical mountain rainforest of Southern Ecuador, were identified by sequencing the whole ITS1-5.8S-ITS2 region and part of 28S rDNA. Mycorrhizae of 13 orchid individuals were investigated by transmission electron microscopy. Simple septal pores and symplechosomes in the hyphal coils of mycorrhizae from four orchid individuals indicated members of Atractiellomycetes. Molecular phylogeny of sequences from mycobionts of 32 orchid individuals out of 103 samples confirmed Atractiellomycetes and the placement in Pucciniomycotina, previously known to comprise only parasitic and saprophytic fungi. Thus, our finding reveals these fungi, frequently associated to neotropical orchids, as the most basal living basidiomycetes involved in mycorrhizal associations of land plants.     

Land‐use History Affects the Distribution of the Saprophytic Orchid Wullschlaegelia calcarata in Puerto Rico's Tabonuco Forest1

Human activities such as logging and agriculture can severely damage forest ecosystems by changing forest structure, ecosystem function, and biodiversity. These changes may have long‐lasting consequences, which influence forest recovery. We investigated the effect of past human disturbance on the current distribution of an understory, achlorophyllous orchid, Wullschlaegelia calcarata in Puerto Rico's tropical rain forest after 70 yr of recovery. Our study site was the 16‐ha Luquillo Forest Dynamics Plot located in the Luquillo Experimental Forest, which has four areas with differing intensity of land use that have been distinguished from variation in canopy cover seen in aerial photographs taken in 1936. We recorded orchids in six 10‐m‐wide, 500‐m‐long transects across four different areas of land‐use history. We found that the orchid was not present in an area of the plot which had <20 percent canopy cover in 1936, and was most abundant in the area with >80 percent canopy cover, which had been minimally impacted by human activity. Tree species composition varied among land‐use history areas, and our observations suggested that this variation might be influencing the local distribution of W. calcarata. We also measured leaf litter biomass and identified the leaves of litter in areas with and without the orchid. Litter with a high proportion of Buchenavia tetraphylla leaves had more orchids. Even though human disturbance ceased in 1932, land‐use history in the Luquillo Forest still casts a shadow over the distribution of W. calcarata.     

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。     

Research on orchid biology and biotechnology

Orchidaceae constitute one of the largest families of angiosperms. They are one of the most ecological and evolutionary significant plants and have successfully colonized almost every habitat on earth. Because of the significance of plant biology, market needs and the current level of breeding technologies, basic research into orchid biology and the application of biotechnology in the orchid industry are continually endearing scientists to orchids in Taiwan. In this introductory review, we give an overview of the research activities in orchid biology and biotechnology, including the status of genomics, transformation technology, flowering regulation, molecular regulatory mechanisms of floral development, scent production and color presentation. This information will provide a broad scope for study of orchid biology and serve as a starting point for uncovering the mysteries of orchid evolution.     

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.     

Orchid Species Richness along Elevational and Environmental Gradients in Yunnan,China

The family Orchidaceae is not only one of the most diverse families of flowering plants, but also one of the most endangered plant taxa. Therefore, understanding how its species richness varies along geographical and environmental gradients is essential for conservation efforts. However, such knowledge is rarely available, especially on a large scale. We used a database extracted from herbarium records to investigate the relationships between orchid species richness and elevation, and to examine how elevational diversity in Yunnan Province, China, might be explained by mid-domain effect (MDE), species–area relationship (SAR), water–energy dynamics (WED), Rapoport’s Rule, and climatic variables. This particular location was selected because it is one of the primary centers of distribution for orchids. We recorded 691 species that span 127 genera and account for 88.59% of all confirmed orchid species in Yunnan. Species richness, estimated at 200-m intervals along a slope, was closely correlated with elevation, peaking at 1395 to 1723 m. The elevational pattern of orchid richness was considerably shaped by MDE, SAR, WED, and climate. Among those four predictors, climate was the strongest while MDE was the weakest for predicting the elevational pattern of orchid richness. Species richness showed parabolic responses to mean annual temperature (MAT) and mean annual precipitation (MAP), with maximum richness values recorded at 13.7 to 17.7°C for MAT and 1237 to 1414 mm for MAP. Rapoport’s Rule also helped to explain the elevational pattern of species richness in Yunnan, but those influences were not entirely uniform across all methods. These results suggested that the elevational pattern of orchid species richness in Yunnan is collectively shaped by several mechanisms related to geometric constraints, size of the land area, and environments. Because of the dominant role of climate in determining orchid richness, our findings may contribute to a better understanding of the potential effects of climate change on orchid diversity, and the development of conservation strategies for orchids.     

Mycorrhizal Fungi Promote Growth and Nitrogen Utilization by Dendrobium nobile (Orchidaceae)

Mycorrhizal associations play a key role in the life cycle and evolutionary history of orchids. Although most orchid species are tropical and epiphytic, their mycorrhizae are poorly understood compared with those of temperate, terrestrial orchids. To investigate the influences of such fungi on photosynthetic, epiphytic orchids, we inoculated seedlings of Dendrobium nobile with Epulorhiza sp. (S1) or Tulasnella sp. (S3). These fungi had been identified based on their morphological and molecular characters. Both S1 and S3 formed symbiotic associations with our seedlings, promoting their growth and development to various degrees. Results from signature experiments with the 15N stable isotope suggested that the utilization of organic nitrogen by orchid seedlings was significantly improved by S1, but not by S3. Dendrobine contents were significantly higher in all inoculated seedlings. Our findings demonstrate that these mycorrhizal fungi enhance plant growth, their utilization of organic nitrogen, and the accumulation of secondary metabolites in this epiphytic orchid species.     

中国1,334种兰科植物就地保护状况评价

作者通过对中国543个自然保护区内兰科植物数据的分析,对我国1,334种兰科植物的就地保护状况进行了逐一评价。我们根据有记录分布保护区的数量,通过专家咨询的方式,将兰科植物就地保护水平划分为"有效保护"、"较好保护"、"一般保护"、"较少保护"、"保护状况不明"和"未予评价"6个等级。结果发现:有676种兰科植物在自然保护区内得到了不同程度的就地保护,占所评价兰科植物总数的51.9%;但有626种兰科植物保护状况不明,尚未受到就地保护,占总数的46.9%。另外有32种仅分布在香港、澳门和台湾地区,本研究中未予评价,约占总数的2.4%。根据评价结果,就兰科植物就地保护中存在的基础调查不足、受关注程度低、受保护率低等问题进行了深入的讨论,提出5条有针对性的建议对策:(1)加强兰科植物的调查监测;(2)提高兰科植物的保护级别与受关注程度;(3)进一步完善兰科自然保护区建设;(4)规范自然保护区综合科学考察;(5)定期评价我国兰科植物保护成效。