Herbaceous vegetation under temperate climatic conditions generally shows nitrogen- or phosphorus-limited plant growth, which creates conditions for a high botanical diversity. Is nitrogen (N) or phosphorus (P) the most important limiting factor, or are both generally in short supply? What are the consequences of Increased N and P inputs that result from anthropogenic disturbances? A new indicator of N or P limitation, that is, the vegetation N:P ratio, allows us to address these questions for a range of mires, dune slacks and moist grasslands.

Nitrogen-or phosphorus-limited growth in herbaceous, wet vegetation : relations with atmospheric input and management regimes

Germination and seedling establishment in orchids: a complex of requirements

� Background Seedling recruitment is essential to the sustainability of any plant population. Due to the minute nature of seeds and early-stage seedlings, orchid germination in situ was for a long time practically impossible to observe, creating an obstacle towards understanding seedling site requirements and fluctuations in orchid populations. The introduction of seed packet techniques for sowing and retrieval in natural sites has brought with it important insights, but many aspects of orchid seed and germination biology remain largely unexplored. � Key Considerations The germination niche for orchids is extremely complex, because it is defined by requirements not only for seed lodging and germination, but also for presence of a fungal host and its substrate. A mycobiont that the seedling can parasitize is considered an essential element, and a great diversity of Basidiomycota and Ascomycota have now been identified for their role in orchid seed germination, with fungi identifiable as imperfect Rhizoctonia species predominating. Specificity patterns vary from orchid species employing a single fungal lineage to species associating individually with a limited selection of distantly related fungi. A suitable organic carbon source for the mycobiont constitutes another key requirement. Orchid germination also relies on factors that generally influence the success of plant seeds, both abiotic, such as light/shade, moisture, substrate chemistry and texture, and biotic, such as competitors and antagonists. Complexity is furthermore increased when these factors influence seeds/seedling, fungi and fungal substrate differentially. � Conclusions A better understanding of germination and seedling establishment is needed for conservation of orchid populations. Due to the obligate association with a mycobiont, the germination niches in orchid species are extremely complex and varied. Microsites suitable for germination can be small and transient, and direct observation is difficult. An experimental approach using several levels of environmental manipulation/control is recommended.

VIEWPOINT: PART OF A HIGHLIGHT ON ORCHID BIOLOGY Germination and seedling establishment in orchids: a complex of requirements

Understanding temporal effects of fire frequency on plant species diversity and vegetation structure is critical for managing tallgrass prairie (TGP), which occupies a mid-continental longitudinal precipitation and productivity gradient. Eastern TGP has contributed little information toward understanding whether vegetation–fire interactions are uniform or change across this biome. We resampled 34 fire-managed mid- and late-successional ungrazed TGP remnants occurring across a dry to wet-mesic moisture gradient in the Chicago region of Illinois, USA. We compared hypotheses that burning acts either as a stabilizing force or causes change in diversity and structure, depending upon fire frequency and successional stage. Based on western TGP, we expected a unimodal species richness distribution across a cover–productivity gradient, variable functional group responses to fire frequency, and a negative relationship between fire frequency and species richness. Species diversity was unimodal across the cover gradient and was more strongly humpbacked in stands with greater fire frequency. In support of a stabilizing hypothesis, temporal similarity of late-successional vegetation had a logarithmic relationship with increasing fire frequency, while richness and evenness remained stable. Temporal similarity within mid-successional stands was not correlated with fire frequency, while richness increased and evenness decreased over time. Functional group responses to fire frequency were variable. Summer forb richness increased under high fire frequency, while C4 grasses, spring forbs, and nitrogen-fixing species decreased with fire exclusion. On mesic and wet-mesic sites, vegetation structure measured by the ratio of woody to graminoid species was negatively correlated with abundance of forbs and with fire frequency. Our findings that species richness responds unimodally to an environmental-productivity gradient, and that fire exclusion increases woody vegetation and leads to loss of C4 and N-fixing species, suggest that these processes are uniform across the TGP biome and not affected by its rainfall–productivity gradient. However, increasing fire frequency in eastern TGP appears to increase richness of summer forbs and stabilize late-successional vegetation in the absence of grazing, and these processes may differ across the longitudinal axis of TGP. Managing species diversity in ungrazed eastern TGP may be dependent upon high fire frequency that removes woody vegetation and prevents biomass accumulation.

Repeated burning of eastern tallgrass prairie increases richness and diversity, stabilizing late successional vegetation

This review summarises scientific knowledge concerning the mycorrhizal associations, pollination, demographics, genetics and evolution of Australian terrestrial orchids relevant to conservation. The orchid family is highly diverse in Western Australia (WA), with over 400 recognised taxa of which 76 are Declared Rare or Priority Flora. Major threats to rare orchids in WA include habitat loss, salinity, feral animals and drought. These threats require science-based recovery actions resulting from collaborations between universities, government agencies and community groups. Fungal identification by DNA-based methods in combination with compatibility testing by germination assays has revealed a complex picture of orchid–fungus diversity and specificity. The majority of rare and common WA orchids studied have highly specific mycorrhizal associations with fungi in the Rhizoctonia alliance, but some associate with a wider diversity of fungi. These fungi may be a key factor influencing the distribution of orchids and their presence can be tested by orchid seed bait bioassays. These bioassays show that mycorrhizal fungi are concentrated in coarse organic matter that may be depleted in some habitats (e.g. by frequent fire). Mycorrhizal fungi also allow efficient propagation of terrestrial orchids for reintroduction into natural habitats and for bioassays to test habitat quality. Four categories of WA orchids are defined by the following pollination strategies: (i) nectar-producing flowers with diverse pollinators, (ii) non-rewarding flowers that mimic other plants, (iii) winter-flowering orchids that attract fungus-feeding insects and (iv) sexually deceptive orchids with relatively specific pollinators. An exceptionally high proportion of WA orchids have specific insect pollinators. Bioassays testing orchid-pollinator specificity can define habitats and separate closely related species. Other research has revealed the chemical basis for insect attraction to orchids and the ecological consequences of deceptive pollination. Genetic studies have revealed that the structure of orchid populations is influenced by pollination, seed dispersal, reproductive isolation and hybridisation. Long-term demographic studies determine the viability of orchid populations, estimate rates of transition between seedling, flowering, non-flowering and dormant states and reveal factors, such as grazing and competition, that result in declining populations. It is difficult to define potential new habitats for rare orchids because of their specific relationships with fungi and insects. An understanding of all three dimensions of orchid habitat requirements can be provided by bioassays with seed baits for fungi, flowers for insects and transplanted seedlings for orchid demography. The majority of both rare and common WA orchids have highly specific associations with pollinating insects and mycorrhizal fungi, suggesting that evolution has favoured increasing specificity in these relationships in the ancient landscapes of WA.

Scientific approaches to Australian temperate terrestrial orchid conservation

Pattern and zonation of peatland vegetation are regulated by environmental gradients, as well as by effects of biomass and competitive exclusion on distribution of species richness. The interplay of these factors has not been closely examined in calcareous prairie fens, which are isolated, species rich, calcareous peatlands in the Prairie Peninsula region of North America. We used multivariate analyses to classify vegetation and to quantify species richness in relation to substrate conditions and vegetation structure in a 23-ha calcareous prairie fen in northeast Illinois, USA. Plant assemblages formed a floristic continuum across sedge meadow, graminoid fen, calcareous seep, marl flat, and spring run vegetation, with complete dissimilarity between spring run and sedge meadow. These vegetation zones corresponded to gradients of decreasing organic content and cation exchange capacity, and increasing pH, Na, Mg, and total Ca concentrations, which reach extremes in spring run and marl flats. Species richness was unimodal across the fen gradient, fitting an expected model of low richness in vegetation either with large biomass (as shown by low light penetration in tall sedge meadow) or with environmentally harsh conditions and low biomass (shown by high light penetration in short marl flat and spring run vegetation). These biotic and abiotic factors, as well as hydrology, mediate vegetation pattern across the fen.

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Relationships among environmental factors, vegetation zones, and species richness in a north American calcareous prairie fen

The Federal threatened eastern prairie fringed orchid (Platanthera leucophaea) occupies prairies, sedge meadows, bogs and fens, primarily north of the Wisconsinan glacial boundary. In the Midwest, where restoration is a recovery objective, its southern distribution is thought to be limited by the transition from nutrient-rich Wisconsinan-aged soils to more acidic nutrient poor soils of Illinoian-aged glacial drift. To better understand edaphic factors affecting its distribution and potential for establishment of new populations, we analyzed soil characteristics across the range of habitats occupied by this species, as well as from unoccupied habitats on the Illinoian Till Plain. We found that P. leucophaea occupies a complex edaphic gradient in variation of % organic matter, base content and soil texture. On Wisconsinan-aged substrates, it occurs in circum-neutral base-rich organic prairie soils in Illinois and Wisconsin and in less calcareous soils with slightly higher pH and lower organic matter content in Michigan lake plain prairies. Eastern sand prairie and sedge meadow habitats on Wisconsinan-aged drift and on unglaciated soils are moderately acid and nutrient poor, while bog and fen habitats are more strongly acidic and highly organic, with no evidence for an underlying calcareous substrate. In comparison, unoccupied prairie soils on the Illinoian till plain have lower pH, % organic matter and base concentrations. These soils also have relatively high % silt content which results in comparatively low available soil moisture holding capacity. This combination of soil conditions may exceed the tolerance limits of P. leucophaea and prevent this species from occurring south of the Wisconsin glacial boundary in the Midwest. On the other extreme, calcareous fens have high pH levels as well as extremely high calcium concentrations, which may exceed the tolerance limits of this species. These findings have implications for guiding efforts to establish P. leucophaea into habitats that should be suitable for this species.

Relationships Between Soil Characteristics, Distribution and Restoration Potential of The Federal Threatened Eastern Prairie Fringed Orchid, Platanthera leucophaea (Nutt.) Lindl

Patrick D. Kelsey

Papers

Relationships among environmental factors, vegetation zones, and species richness in a north American calcareous prairie fen

Relationships Between Soil Characteristics, Distribution and Restoration Potential of The Federal Threatened Eastern Prairie Fringed Orchid, Platanthera leucophaea (Nutt.) Lindl