Roadsides provide refuge for orchids: characteristic of the surrounding landscape

Seminatural habitats are declining throughout the world; thus, the role of small anthropogenic habitats in the preservation of plants is becoming increasingly appreciated. Here, we surveyed the orchid flora of roadside verges in five Central European countries (Austria, Hungary, Romania, Slovakia, and Slovenia) and tested how the surrounding landscape matrix affects the overall number of species and individuals, and also different functional groups of orchids. We found more than 2,000 individuals of 27 orchid species during our surveys. According to our results, the increasing coverage of agricultural and urban areas negatively affects both the number of orchid species and individuals on roadsides. Our study further suggests that differences in the surrounding habitats affect which species are found on roadsides, since the increasing coverage of grasslands or forested areas around orchid occurrences had a significant positive effect on the number of grassland or forest‐dwelling species and individuals, respectively. Most variance in orchid numerosity and diversity was explained by the cover of the suitable habitat types of the respective taxa in the surrounding landscape of the sampling points. This highlights the importance of roadsides acting as refugia for numerous species and valuable plant communities as well as in supporting biodiversity in general.     

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.     

Consistency in the habitat degree of invasion for three invasive plant species across Mediterranean islands

Habitats are known to vary in their vulnerability to invasion by alien plants and different species often colonise distinct habitats. To assess the consistency in the degree of invasion of particular habitats, this study examined the frequency of occurrence and local abundance of three invasive plant taxa: Ailanthus altissima, Carpobrotus spp., and Oxalis pes-caprae across different habitat types on four representative Mediterranean islands. We conducted systematic field surveys recording the presence-absence and cover of these taxa on the islands of Mallorca, Corsica, Sardinia and Crete. Drawing on the results of 5,285 sample points, the frequency of occurrence of the three invaders tends to be higher than expected in urban, ruderal and roadside habitats. In contrast, scrub habitats rarely contain any of the three invaders, indicating that they may be more resistant to invasion. The degree of invasion, determined by the local abundance of an invasive plant in any one habitat, varies according to the identity of the invader and the island. However, based on average abundance, Oxalis pes-caprae exhibits the highest degree of invasion, and Carpobrotus spp. the least. There is no indication that any one of the four islands is more prone to either higher frequencies or abundances of the three invaders. These patterns suggest that anthropogenic changes in Mediterranean islands will increase the vulnerability of certain habitats to invasion and increase the distribution of these three invasive taxa at any of the four islands.     

Living on the verge: are roads a more suitable refuge for small mammals than streams in Mediterranean pastureland?

The retention of natural habitat corridors is a useful and practical conservation tool that can attenuate the effects of habitat loss and fragmentation on wildlife. Linear structures may contribute to the conservation of biodiversity by providing additional habitats for small fauna living in highly modified environments. We assessed the importance of road verges as refuge areas for small mammals, in highly intensified grazed pastures, within a Mediterranean landscape and compared the role of road verges as refuges with that of riparian galleries, which have been described as important shelter locations for small fauna. For this purpose, a small mammal trapping study was undertaken on two road verges and beside two small streams in southern Portugal. We captured 457 individuals of five different species, with Mus spretus the most common species captured, followed by Crocidura russula. Captures were 4.6-fold higher immediately beside both roads and streams than 12 m away in the surrounding matrix. Individuals captured in the matrix presented a smaller body size and lower body condition, suggesting that this suboptimal habitat is occupied mainly by subadults. M. spretus was 46% more abundant by roads than by streams, while C. russula was present in similar numbers in both habitats. M. spretus individuals were larger near streams but exhibited no difference in body condition between habitats. C. russula had a better body condition and slightly higher body lengths at roadsides. Our results show that roadside verges in intensively grazed Mediterranean landscapes act as important refuges and constitute equally vital habitats for small mammals as do riparian vegetation strips in landscapes where other suitable habitats are scarce.     

Pollination Syndromes in Mediterranean Orchids—Implications for Speciation,Taxonomy and Conservation

The Mediterranean flora is spectacularly rich in orchid species that have evolved remarkable adaptations to their environment. Orchids have complex and delicate interactions with their pollinators, which makes them particularly prone to local extinction. Conservation actions should be encouraged for a range of endangered Mediterranean orchid species, but the current taxonomic confusion in several genera and the apparent disagreement among orchid taxonomists make the situation particularly confusing from a conservation perspective. In this review, we document how the different pollination syndromes of Mediterranean orchids (nectar reward, shelter offering, food deception and sexual deception) can have a profound impact on the type of reproductive barriers among species, on floral phenotypic variation as we perceive it, on potentially related processes of species sorting and extinction and, consequently, should have a strong influence on the related conservation management programs. We also highlight that the majority of Mediterranean orchids are pollinated by specialised bees often occupying otherwise narrow ecological niches (e.g. pollen specialisation, brood cell parasites, specific nesting site). This condition makes the orchid-pollinator interactions very fragile and several orchid species prone to local extinction. We illustrate this phenomenon by a selection of case studies that show how the adequate integration of the ecological requirements/traits of the orchids and their associated pollinators into conservation actions could help protect endangered species and ensure the sustainability of the often complex local pollination web.     

Turkish graveyards as refuges for orchids against tuber harvest

Harvest of orchid tubers for salep production is widespread in southwestern Asia and the Balkans and constitutes a major conservation risk for wild orchid populations. Synanthropic habitats, such as graveyards, are important refuges for orchids and other organisms and could offer protection from salep harvesting because of their special cultural role. However, little is known about the occurrence and factors influencing harvesting of salep in graveyards. During field surveys of 474 graveyards throughout Turkey, we observed 333 graveyards with orchids, 311 graveyards with tuberous orchids, and salep harvest in 14 graveyards. Altogether, 530 individuals of 17 orchid species were collected, representing 9% of the individuals recorded. Harvesting intensity was relatively low, and populations were usually not wholly destroyed. However, some species were clearly more affected than others. Salep harvesting risk of orchid species was significantly associated with flowering time, with early‐flowering species being more affected. A marginally significant positive relationship between harvesting risk and species‐specific tuber size was also detected. Our data suggest that graveyards might offer some protection against salep harvesting in Turkey, but they also show that some orchid taxa are much more affected than others. Overall, our observations add more weight to the conservation value of these special habitats.     

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.     

Butterfly diversity in Mediterranean islands and in Pentadaktylos <Emphasis Type="Italic">Pinus</Emphasis><Emphasis Type="BoldItalic"> </Emphasis><Emphasis Type="Italic">brutia</Emphasis> forests of Cyprus

We analysed the influence of contemporary geography on butterfly diversity for islands in the Mediterranean Basin. We found that island size and distance from the mainland has a significant effect on the number of species. We also used butterflies as an indicator group to identify the importance of forest habitats for biodiversity conservation in the island of Cyprus. To understand the relative importance of local vegetation characteristics of butterflies in the Pentadaktylos mountains transect counts were used to assess the abundance and butterfly diversity in two different forest types. A total of 1,602 butterflies and 23 species were recorded during this research. We observed highly significant effects of forest type on abundance and species richness of butterflies. For example, number of butterflies was significantly higher in old forest than young pine forest. Also, the abundance of endemic butterflies was highest in old forest habitats. Therefore, the survival of the majority of endemic butterflies in Cyprus may depend on conservation of old forests and their understorey plants.     

Relationships between orchid and fungal biodiversity: Mycorrhizal preferences in Mediterranean orchids

Orchidaceae is one of the most species-rich angiosperm families, and all orchids are fully dependent on fungi for their seed germination and their life cycle. The level of specificity of the association between orchid species and fungi can be related to the number of co-occurring orchid species. To investigate orchid mycorrhizal associations in adult-photosynthetic orchids, 16 Mediterranean orchid species belonging to 4 genera (Anacamptis, Ophrys, Orchis, and Serapias) at 11 different sites were subjected to DNA-based analysis. Eighteen operational taxonomic units representing two fungal families, Tulasnellaceae and Ceratobasidiaceae, were identified. All examined orchid species associated with different mycorrhizal fungi. Interestingly, there was a positive correlation between number of orchid species and number of mycorrhizal. Monospecific populations showed a lower number of fungi, while sympatric populations had a higher number of mycorrhizal fungi. Our results showed that Mediterranean orchid species associated with a higher number of mycorrhizal fungi confirming as photosynthetic orchids are typically generalists toward mycorrhizal fungi. Thus, photosynthetic orchids exhibit low specificity for fungal symbionts showing the potential for opportunistic associations with diverse fungi reducing competition for nutrient. We suggest that these characteristics could confer symbiotic assurance particularly in habitat with resource limitations or prone to stressful conditions.     

Conservation value of roadside verges for stenotopic heathland Carabidae: corridors or refugia?

It has been shown that, in The Netherlands, sandy roadside verges may aid the survival of stenotopic Carabidae associated with open habitats such as heathland and sandy grassland. It is suggested that roadsides serve as corridors linking heathland fragments, thus maintaining a metapopulation structure, which helps avoid regional extinction. However, a counter-argument is that verges encourage dispersal into unsuitable habitats, and function as sinks in which individuals die without reproducing. The stenotopic species recorded from roadside verges but not in adjacent open habitats all had high powers of dispersal. The least successful dispersers were found only in open habitats and in verges closely connected to them. From this, the conclusion that verges function as corridors can be drawn. We reinterpret these data to suggest that stenotopic species were breeding on the roadside verges, and that they acted as refugia rather than corridors. We also present evidence from a roadside verge in the Breckland region of East Anglia, England. We show that many stenotopic species breed on the verge, which thus functions as a refugium. For Carabidae dependent on early successional habitats, survival may be more likely on disturbed roadside verges than on nature reserves in the absence of regular management disturbance.     

Conservation of the Native Orchids Through Seedling Culture and Reintroduction—A Singapore Experience

Singapore is located near the equator, off the southern tip of the Malay Peninsula. The whole country consists mostly of lowland. It has many interesting types of natural habitats such as primary rain forest, freshwater swamp forest, mangroves, secondary forests, shrub, grasslands, and urban parks and fields. The climate is equatorial with relatively uniform temperature and high humidity. Unfortunately, many of the natural habitats and the native orchids which thrive there have disappeared due to habitat destruction. Some 226 species of native orchids have been recorded in Singapore. However, of these 178 are considered to be extinct, and only five are common. The orchid conservation programme aims to monitor existing species, explore ways to conserve their germplasm, and increase their numbers in natural, semi-natural, and urban environments through ex-situ seedling culture and subsequent re-introduction into appropriate habitats, including roadside trees, parks and natural areas. In the first phase of the programme, we have successfully propagated and carried out experiments of re-introduction on five species of native orchids, namely, Grammatophyllum speciosum, Bulbophyllum vaginatum, Bulbophyllum membranaceum, Cymbidium finlaysonianum and Cymbidium bicolor. Survival percentages 8-yr after the reintroduction events ranged from 10 to 95 for G. speciosum, the target species of the earliest re-introduction experiments. Size of the seedlings at reintroduction, host trees, and relative humidity seemed to play significant roles in the success rate of the reintroductions.     

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.     

Roadside verges can support spontaneous establishment of steppe-like habitats hosting diverse assemblages of bees and wasps (Hymenoptera: Aculeata) in an intensively cultivated central European landscape

Anthropogenic habitats, which are frequently occupied by otherwise endangered species of the open landscape, suffer from a lack of sustainability. Here, we analyzed one of the few sustainable anthropogenic habitats, roadside verges. We hypothesized that steppe-like habitats at upper parts of south-exposed slopes of roadside verges may serve as the only strongholds of threatened species ousted from the surrounding intensively cultivated landscape, with direct consequences for plant pollination and other ecosystem services. We examined bees and wasps alongside Highway 7 constructed 14–28 years ago in the northwestern Czech Republic. The sampling sites were located 4–17 m from paved road with traffic intensity of 9523 to 38,688 vehicles × 24 h^?1, and they consisted of steppe-like habitats with patches of bare soil (sand, loess and claystone) that formed and were maintained spontaneously. The sites were not connected to any permanent grasslands or forests, and they were surrounded mostly by arable fields. We found 139 species of herbs (nine threatened) in vegetation surveys along transects. We identified 164 species of bees and wasps, with a dominance of 0.052 and a Fisher’s alpha of 45.0; 32 species were threatened, including a second Czech record of Chrysis gribodoi spilota and four critically endangered species (Andrena florivaga, Halictus scabiosae, Sphecodes spinulosus and Nanoclavelia leucoptera). The captured specimens consisted of a higher number of loess specialists and other steppe species compared to specimens associated with other anthropogenic habitats that provide bare ground (sand pits and ash deposits). In conclusion, the formation of spontaneously maintained steppe-like habitats at roadside verges should be considered to be an appropriate tool supporting the biodiversity in the highly cultivated landscape of central Europe.     

广东兰科植物多样性保育现状

为了使广东省的兰科植物及其遗传多样性得到有效的保育, 保存我国重要野生植物资源, 在2017-2019年间, 采用样线和样地相结合的调查手段、专家快速评估和野外调查相结合的评估技术以及Wilcoxon符号秩检验和Friedman检验的统计方法, 对广东省自然分布的兰科植物进行了全面的调查和濒危等级评估, 并对其在广东省自然保护区中的就地保育情况和全国植物园中的迁地保育情况进行了综合分析。结果表明, 广东省分布有兰科植物80属235种, 其中广东特有种20种; 广东兰科植物受威胁物种有186种, 其中极危11种、濒危114种、易危61种; 就地保育的兰科植物有111种, 迁地保育的兰科植物有156种, 就地和迁地共同保育的兰科植物有96种, 保育的有效程度较低; 另外, 就地、迁地、就地和迁地共同保育的兰科植物之间没有体现出明显的差异, 保育工作缺乏选择性和针对性。基于此, 我们建议广东兰科植物的保育工作应重视基础数据的收集和持续的野外监测、提高保育物种的数量、优化迁地保育物种的选择性和针对性、完善迁地保育和就地保育之间的协同性, 同时也应重视立法和公众教育, 并构建广东兰科植物保育的网络系统。     

Local and regional abundance of exotic plant species on Mediterranean islands: are species traits important?

Aim We assess the importance of three relevant and readily obtainable life‐history traits (dispersal syndrome, stem height and growth form) and biogeographical origin (European vs. non‐European) on the local and regional abundance of over 400 exotic plant species across eight Mediterranean islands. Location The Mediterranean islands of Lesbos, Rhodes, Crete, Malta, Corsica, Sardinia, Majorca and Minorca. Methods We adopt two abundance criteria for each exotic species: the proportion of islands in which the species occurs (regional abundance), and a qualitative estimate of species abundance within each of five islands (local abundance). Subsequently, we assess the relationship between local and regional abundance, as well as the role of key life‐history traits on both regional and local abundance. These analyses were undertaken separately for the European exotics and the non‐European exotics. Results Only 10.9% of the species occur on more than four islands, and only four species are present on all eight islands. Both local and regional abundances were higher for the non‐European than the European species. Local and regional abundances were positively correlated, particularly for exotics with non‐European origins. Wind‐dispersed species tended to have higher regional abundance than species dispersed by other means but this trend only occurred for local abundance on two islands — Corsica and Majorca. Neither a species’ growth form nor its stem height explained trends in regional or local abundance. Conclusions Although wind‐dispersed exotics are more widespread in the Mediterranean, plant life‐history traits appear to play a lesser role in invasion success than area of biogeographical origin. In general, exotic species of non‐European origin were more abundant at both local and regional scales. Invasion patterns should be interpreted at both local and regional scales, but the stochastic nature of biological invasions may limit deterministic interpretations of invasion patterns, especially if islands are studied in isolation.     

Evidence for pollinator sharing in Mediterranean nectar-mimic orchids: absence of premating barriers?

Pollinator specificity has traditionally been considered the main reproductive isolation mechanism in orchids. Among Mediterranean orchids, however, many species attract and deceive pollinators by mimicking nectar-rewarding plants. To test the extent to which deceptive orchid species share pollinators, we collected and identified hemipollinaria-carrying insects, and used ribosomal sequences to identify the orchid species from which hemipollinaria were removed. We found that social and solitary bees, and also flies, carried hemipollinaria belonging to nine orchid species with different degrees of specialization. In particular, Anacamptis morio, Dactylorhiza romana and Orchis mascula used a large set of pollinator species, whereas others such as Orchis quadripunctata seemed to be pollinated by one pollinator species only. Out of the insects with hemipollinaria, 19% were found to carry hemipollinaria from more than one orchid species, indicating that sympatric food-deceptive orchids can share pollinators. This sharing was apparent even among orchid sister-species, thus revealing an effective overlap in pollinator sets among closely related species. These results suggest varying degrees of pollinator specificity in these orchids, and indicate that pollinator specificity cannot always act as the main isolation mechanism in food-deceptive temperate orchids.     

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。     

Bird communities of highway verges: Influence of adjacent habitat and roadside management

We have investigated the effects of landscape traversed and roadside structure on the use of highway verges by birds. Three contrasted landscapes were chosen in terms of human land use and vegetation structure: an intensive farmland, a pine plantation, and a matoral. The roadside sections varied in vegetation structure, width and profile. We recorded birds present in roadsides and adjacent habitats by transect counts over all seasons. Roadside bird species appeared for a great part similar to those of adjacent habitats. However, diversity and abundance in verges did not depend on that of adjacent habitats. Woody roadsides were comparable to hedges, as trees (and shrubs) in verges enhanced species richness and abundance of birds in the farmland and woodland sites. Width and profile of verges had less influence. In all sites, typical species of the habitat traversed partly avoided roadsides. On the contrary, numerous species associated with ‘rare’ habitats in one site preferred roadsides, provided that verge vegetation contrasted with the dominant habitat. It is concluded that birds responses to highways can vary greatly with landscape traversed and verge vegetation. Highway verges could be favorable to birds, if they constitute a complementary habitat to the dominant habitat within a landscape.     

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.     

A narrowly endemic photosynthetic orchid is non‐specific in its mycorrhizal associations

Mycorrhizal association is a common characteristic in a majority of land plants, and the survival and distribution of a species can depend on the distribution of suitable fungi in its habitat. Orchidaceae is one of the most species‐rich angiosperm families, and all orchids are fully dependent on fungi for their seed germination and some also for subsequent growth and survival. Given this obligate dependence, at least in the early growth stages, elucidating the patterns of orchid–mycorrhizal relationships is critical to orchid biology, ecology and conservation. To assess whether rarity of an orchid is determined by its specificity towards its fungal hosts, we studied the spatial and temporal variability in the host fungi associated with one of the rarest North American terrestrial orchids, Piperia yadonii. The fungal internal transcribed spacer region was amplified and sequenced by sampling roots from eight populations of P. yadonii distributed across two habitats, Pinus radiata forest and maritime chaparral, in California. Across populations and sampling years, 26 operational taxonomic units representing three fungal families, the Ceratobasidiaceae, Sebacinaceae and Tulasnellaceae, were identified. Fungi belonging to the Sebacinaceae were documented in orchid roots only at P. radiata forest sites, while those from the Ceratobasidiaceae and Tulasnellaceae occurred in both habitats. Our results indicate that orchid rarity can be unrelated to the breadth of mycorrhizal associations. Our data also show that the dominance of various fungal families in mycorrhizal plants can be influenced by habitat preferences of mycorrhizal partners.