Drivers of vegetative dormancy across herbaceous perennial plant species.

Vegetative dormancy, that is the temporary absence of aboveground growth for ≥ 1 year, is paradoxical, because plants cannot photosynthesise or flower during dormant periods. We test ecological and evolutionary hypotheses for its widespread persistence. We show that dormancy has evolved numerous times. Most species displaying dormancy exhibit life-history costs of sprouting, and of dormancy. Short-lived and mycoheterotrophic species have higher proportions of dormant plants than long-lived species and species with other nutritional modes. Foliage loss is associated with higher future dormancy levels, suggesting that carbon limitation promotes dormancy. Maximum dormancy duration is shorter under higher precipitation and at higher latitudes, the latter suggesting an important role for competition or herbivory. Study length affects estimates of some demographic parameters. Our results identify life historical and environmental drivers of dormancy. We also highlight the evolutionary importance of the little understood costs of sprouting and growth, latitudinal stress gradients and mixed nutritional modes.

Genetic structure of a regionally endangered orchid, the dark red helleborine (Epipactis atrorubens) at the edge of its distribution.

The genetic structure and diversity of species is determined by both current population dynamics and historical processes. Population genetic structure at the edge of the distribution is often expected to differ substantially from populations at the centre, as these edge populations are often small and fragmented. In addition, populations located in regions that have experienced repeated glaciations throughout the Pleistocene, may still carry imprints from the genetic consequences of frequent distribution shifts. Using chloroplast DNA sequences and nuclear microsatellite markers we studied the genetic structure of Epipactis atrorubens at the northern edge of its distribution. Contrary to populations in the centre of the distribution, populations at the northern range are regionally endangered as they are small and disjunct. Sequence data of 2 chloroplast loci and allelic data from 6 nuclear microsatellite markers were obtained from 297 samples from Finland, Estonia and Russia. We sought for genetic indicators of past population processes, such as post-glacial colonisation history of E. atrorubens. As expected, we observed low genetic variation, in terms of numbers of substitutions, haplotypes and alleles, and significant levels of differentiation, especially pronounced in the chloroplast DNA. These features suggest that the edge populations could be prone to extinction.

Factors influencing IUCN threat levels to orchids across Europe on the basis of national red lists.

The red list has become a ubiquitous tool in the conservation of species. We analyzed contemporary trends in the threat levels of European orchids, in total 166 species characterized in 27 national red lists, in relation to their reproductive biology and growth form, distribution area, and land cover where they occur. We found that species in central Europe are more threatened than those in the northern, southern, or Atlantic parts of Europe, while species were least threatened in southern Europe. Nectarless and tuberous species are significantly more threatened than nectariferous and rhizomatous taxa. Land cover (ratios of artificial land cover, area of pastures and grasslands, forests and inland wetlands) also significantly impacted the threat level. A bigger share of artificial land cover increases threat, and a bigger share of pasture and grassland lowers it. Unexpectedly, a bigger share of inland wetland area in a country increased threat level, which we believe may be due to the threatened nature of wetlands themselves relative to other natural land cover types. Finally, species occurring in multiple countries are on average less threatened. We believe that large-scale analysis of current IUCN national red lists as based on their specific categories and criteria may particularly inform the development of coordinated regional or larger-scale management strategies. In this case, we advocate for a coordinated EU protection and restoration strategy particularly aimed at central European orchids and those occurring in wetland area.

Minority cytotypes in European populations of the Gymnadenia conopsea complex (Orchidaceae) greatly increase intraspecific and intrapopulation diversity.

BACKGROUND AND AIMS: Patterns of ploidy variation among and within populations can provide valuable insights into the evolutionary mechanisms shaping the dynamics of plant systems showing ploidy diversity. Whereas data on majority ploidies are, by definition, often sufficiently extensive, much less is known about the incidence and evolutionary role of minority cytotypes. METHODS: Ploidy and proportions of endoreplicated genome were determined using DAPI (4',6-diamidino-2-phenylindole) flow cytometry in 6150 Gymnadenia plants (fragrant orchids) collected from 141 populations in 17 European countries. All widely recognized European species, and several taxa of less certain taxonomic status were sampled within Gymnadenia conopsea sensu lato. KEY RESULTS: Most Gymnadenia populations were taxonomically and/or ploidy heterogeneous. Two majority (2x and 4x) and three minority (3x, 5x and 6x) cytotypes were identified. Evolution largely proceeded at the diploid level, whereas tetraploids were much more geographically and taxonomically restricted. Although minority ploidies constituted <2 % of the individuals sampled, they were found in 35 % of populations across the entire area investigated. The amount of nuclear DNA, together with the level of progressively partial endoreplication, separated all Gymnadenia species currently widely recognized in Europe. CONCLUSIONS: Despite their low frequency, minority cytotypes substantially increase intraspecific and intrapopulation ploidy diversity estimates for fragrant orchids. The cytogenetic structure of Gymnadenia populations is remarkably dynamic and shaped by multiple evolutionary mechanisms, including both the ongoing production of unreduced gametes and heteroploid hybridization. Overall, it is likely that the level of ploidy heterogeneity experienced by most plant species/populations is currently underestimated; intensive sampling is necessary to obtain a holistic picture.

Advantages of volunteer-based biodiversity monitoring in Europe.

Without robust and unbiased systems for monitoring, changes in natural systems will remain enigmatic for policy makers, leaving them without a clear idea of the consequences of any environmental policies they might adopt. Generally, biodiversity-monitoring activities are not integrated or evaluated across any large geographic region. The EuMon project conducted the first large-scale evaluation of monitoring practices in Europe through an on-line questionnaire and is reporting on the results of this survey. In September 2007 the EuMon project had documented 395 monitoring schemes for species, which represents a total annual cost of about 4 million euro, involving more than 46,000 persons devoting over 148,000 person-days/year to biodiversity-monitoring activities. Here we focused on the analysis of variations of monitoring practices across a set of taxonomic groups (birds, amphibians and reptiles, mammals, butterflies, plants, and other insects) and across 5 European countries (France, Germany, Hungary, Lithuania, and Poland). Our results suggest that the overall sampling effort of a scheme is linked with the proportion of volunteers involved in that scheme. Because precision is a function of the number of monitored sites and the number of sites is maximized by volunteer involvement, our results do not support the common belief that volunteer-based schemes are too noisy to be informative. Just the opposite, we believe volunteer-based schemes provide relatively reliable data, with state-of-the-art survey designs or data-analysis methods, and consequently can yield unbiased results. Quality of data collected by volunteers is more likely determined by survey design, analytical methodology, and communication skills within the schemes rather than by volunteer involvement per se.

Mycorrhizal interactions of orchids colonizing Estonian mine tailings hills.

Northeastern Estonia is home to extensive oil shale mines. Associated with these are desolate and environmentally damaging hills of ash and semicoke tailings. Interestingly, some of the first plants to colonize these hills are rare orchids. Here, we assess the identities of the mycorrhizal fungi associated with these orchids, in particular Epipactis atrorubens, Orchis militaris, and Dactylorhiza baltica, and compare them with mycorrhizal fungi from orchids from pristine habitat. Epipactis atrorubens associated with the widest breadth of fungi, including unnamed members of the basidiomycete family Tulasnellaceae and the potentially ectomycorrhizal ascomycetes Trichophaea woolhopeia and Geopora cooperi. Orchis militaris also associated with unnamed members of the Tulasnellaceae. Dactylorhiza baltica associated with Ceratobasidium albasitensis. In Epipactis and Orchis, the same fungi associated with plants in the pristine habitat as with those on ash hills. The tulasnelloid and ceratobasidioid fungi mycorrhizal with these orchids appear closely related to common orchid mycorrhizal fungi, while one of the ascomycetes mycorrhizal with E. atrorubens is closely related to a mycorrhizal fungus with E. microphylla. Our results suggest that these orchids and their fungi are not limited to pristine habitats and that environmentally polluted sites may present novel habitats that may be exploited for endangered plant conservation.

The evolutionary history of mycorrhizal specificity among lady's slipper orchids.

Although coevolution is acknowledged to occur in nature, coevolutionary patterns in symbioses not involving species-to-species relationships are poorly understood. Mycorrhizal plants are thought to be too generalist to coevolve with their symbiotic fungi; yet some plants, including some orchids, exhibit strikingly narrow mycorrhizal specificity. Here, we assess the evolutionary history of mycorrhizal specificity in the lady's slipper orchid genus, Cypripedium. We sampled 90 populations of 15 taxa across three continents, using DNA methods to identify fungal symbionts and quantify mycorrhizal specificity. We assessed phylogenetic relationships among sampled Cypripedium taxa, onto which we mapped mycorrhizal specificity. Cypripedium taxa associated almost exclusively with fungi within family Tulasnellaceae. Ancestral specificity appears to have been narrow, followed by a broadening after the divergence of C. debile. Specificity then narrowed, resulting in strikingly narrow specificity in most of the taxa in this study, with no taxon rewidening to the same extant as basal members of the genus. Sympatric taxa generally associated with different sets of fungi, and most clades of Cypripedium-mycorrhizal fungi were found throughout much of the northern hemisphere, suggesting that these evolutionary patterns in specificity are not the result of biogeographic lack of opportunity to associate with potential partners. Mycorrhizal specificity in genus Cypripedium appears to be an evolvable trait, and associations with particular fungi are phylogenetically conserved.