Factors affecting persistence of terrestrial orchids in wet meadows and implications for their conservation in a changing agricultural landscape

We examined the occurrence of Dactylorhiza majalis, the most abundant terrestrial orchid species growing in rapidly disappearing wet meadows, at 50 historical sites for 3 years. We aimed to find the most frequent reasons for its recent extinction at many sites. We found that the main reasons for its extinction were absence of mowing, intensive fertilisation and washouts of fertilisers from fields nearby. At extant sites, we studied its biometric characteristics and composition of surrounding vegetation, to determine factors affecting its persistence. Bad performance of persisting populations of this species was associated with prevalence of grasses, low May temperatures and absence of mowing. This confirms, at metapopulation level, what has previously been observed at the level of individual populations. We suggest that the system of agricultural subsidies in the country should change towards more sensitive allocation of funds to those farmers, who will adopt the appropriate management of wet meadows and their surroundings.     

Long-term influence of mowing on population dynamics in the rare orchid Dactylorhiza lapponica: The importance of recruitment and seed production

Many orchids are currently red-listed due to changes in land use, and their future persistence will depend on management. Traditional land use like mowing is believed to favour orchids through increased survival and reproduction of established individuals, but the lack of data connecting their complex life cycle presently limits our ability to evaluate effects of management. Here we used data from 16 years of demographic monitoring to study how mowing affects population dynamics in two populations of the rare orchid Dactylorhiza lapponica. Both populations were characterized by long-lived individuals, low adult mortality and high seedling mortality. The traditional regime of mowing every second year strongly increased recruitment and reduced seed production in both populations, but had moderate effect on adult survival, growth rate and flowering probability. Population growth rate was positive for all population × treatment combinations. Traditional mowing significantly increased growth rate in both populations, and LTRE-analyses revealed that this primarily was a result of increased recruitment. The results indicate that demographic rates commonly associated with orchid persistence may be insensitive to traditional management, and underscores the importance of seed production and recruitment to maintain population growth in this long-lived species. The combination of low establishment success and no seed bank makes D. lapponica dependent on high seed input. Our results suggest that D. lapponica would benefit most from traditional mowing performed after seed dispersal in the study areas, but also suggest a high probability of future survival in the absence of mowing.     

The effectiveness of conservation measures to enhance nest survival in a meadow bird suffering from anthropogenic nest loss

The main reason for meadow bird declines is supposed to be the intensification of grassland management with earlier first harvest dates and more frequent harvests, resulting in high nest destruction rates. To increase productivity of meadow bird populations in intensified grassland areas a delay of mowing date and individual nest protection measures have been proposed. However, for ground-nesting songbirds such as the whinchat (Saxicola rubetra) the effectiveness of such measures remains widely unknown. In particular, if nest predation rate is high, measures to protect nests from agricultural destruction alone might be questionable. Here, we quantify whinchat nest survival of (1) unprotected nests situated in early mown meadows, (2) protected nests situated in early mown meadows, and (3) nests situated in late mown meadows. Analyses considered the fact that successful and unsuccessful nests are not found with equal probabilities. Periods of reduced nest survival were associated with mowing periods in the different types of meadows. In early mown meadows nest survival rates were low (S < 0.1), and both conservation measures, individual nest protection and delayed mowing, resulted in significantly increased nest survival rates (S > 0.7). Individual nest protection cannot avoid changes in habitat quality of intensively managed meadows, and therefore is only suitable as small-scale and short-term measure to increase nest success until a high portion of late mown meadows is established. Thus, we suggest that a combination of the two measures applied to intensified grassland fields should be provided to maintain viable sizes of endangered meadow bird populations.     

Local extinctions of the wetland specialist Swertia perennis L. (Gentianaceae) in Switzerland: a revisitation study based on herbarium records

We studied population extinction of the locally abundant fen plant Swertia perennis in Switzerland and used up to 127-year old herbarium records to relocate 63 sites that had once hosted this species. We recorded current site characteristics and related them to the absence or abundance of populations. Fifty-four sites (86%) were still traditionally used (extensively mown or grazed). Fifteen populations (24%) had gone extinct. Extinction was more likely at lower altitude, in the peripheral distribution range of S. perennis (58% peripheral, 9% central populations extinct), on the smallest fens (75% extinct on fens <400 m²) and on fens with intensified land use. However, even on traditionally managed wetlands 18.5% of the populations had gone extinct. Moreover, 40% of all remaining populations were smaller than 250 flowering plants. We conclude that both intensified agricultural practice and habitat fragmentation contributed to local extinction of S. perennis. Small populations, especially, may not be able to persist in the long term.     

Coppice management effects on experimentally established populations of three herbaceous layer woodland species

Traditional coppice management creates a temporal release of resources. We determined how this affected three herbaceous species (Cardamine pratensis, Primula elatior and Geum urbanum) and if it was suitable for their conservation within woodland given their differing phenologies and habitat affinities for woodland.Reproductive adults were transplanted and their fate, i.e. survival and counts of leafs and flowers, plus the fate of their progeny, were monitored for three years following cutting of coppice shoots (three light levels) or yearly autumn mowing.Cardamine pratensis and P. elatior produced more flowers with increasing light availability. Mowing increased flower and leaf production with time for P. elatior. Seedling numbers followed a similar trend. Geum urbanum initially produced more flowers with increasing light and when mown, but the effect disappeared and did not increase seedling numbers. Its basal leaves showed the opposite pattern. Population growth rates (λ), calculated for P. elatior and G. urbanum, confirmed the strong treatment effects on the former and the absence of effects on the latter. Yet, decomposition of treatment effects, showed considerable flexibility in life history of G. urbanum, except for contributions of fecundity. The latter, however, contributed most to positive effects on λ for P. elatior.Early flowering species with an affinity for open habitats (C. pratensis > P. elatior) benefited more from temporal resource release than the later flowering, typical woodland species. Coppice management thereby maintains both typical forest herbs and herbs with affinity for more open habitats. This is an important conservation tool especially in intensively managed landscapes.     

Effect of low-intensity grazing on the species-rich vegetation of traditionally mown subalpine meadows

Subalpine meadows, which traditionally were mown every other year, are particularly rich in plant species, especially forbs. Near Davos (Switzerland) we compared the vegetation of mown sites with that of sites grazed for up to 50 years by non-lactating cows. We recorded an overall mean of 51.5 plant species per 4×4 m plot. Among grazed sites, evenness decreased with time since conversion to grazing (−0.11 in 50 years; P<0.05), suggesting progressive vegetation change, which may eventually result in the loss of species. Ground cover by forbs tended to be higher in mown than in grazed sites (by 7.2%; P<0.1). The proportion of not-clonally colonising perennial species decreased after conversion to grazing (−7.72%; in 50 years, P<0.05), while the cover by graminoid species increased (+14.2% in 50 years; P<0.1). More intensively grazed sites had a lower cover of dwarf shrubs and higher cover of legume species (P<0.05). Because grazing negatively affects both botanical richness and agricultural quality, mowing of traditionally mown subalpine meadows should be maintained, and recently grazed meadows should be reconverted to mowing.     

Future prospects for the rare, late-flowering Gentianella germanica and Gentianopsis ciliata in Dutch nutrient-poor calcareous grasslands

We discuss the population biology of two calcareous grassland gentians, Gentianella germanica and Gentianopsis ciliata, in relation to the habitat management currently practiced in The Netherlands. There, at the margin of their range, both species are rare. Gentianella germanica persists on six remaining locations, whereas Gentianopsis ciliata, with two populations, is nearly extinct. Gentianella germanica is a strict biennial, Gentianopsis ciliata an iteroparous perennial. Both species depend on insects for seed production and suffer from low insect visitation. Pollination experiments in one Dutch population demonstrated that Gentianopsis ciliata is self-compatible, but hardly sets any seed under natural conditions due to pollen limitation. The low reproductive success of both species is partly due to the low pollinator densities at their late flowering time, partly caused by the small population size of the gentians themselves, and partly a result of mowing too early. The latter has destroyed the seed crop of several subsequent years in one population of Gentianopsis ciliata and some of Gentianella germanica. In Gentianella germanica, the early mowing and low insect visitation seems to have resulted in selection of less herkogamous and consequently more autofertile individuals. The perspectives for Gentianopsis ciliata are currently extremely poor in The Netherlands. Under the present circumstances, extinction will most likely occur within 10-20 years. Population reinforcement (seeding, artificial cross-pollination with nearby populations) should be considered if we want to conserve this species. For both gentians, but also of other flagship species of nutrient-poor calcareous grasslands, the total grassland area needs to be enlarged and must constitute an interconnected network of reserves. The traditional management method, sheep grazing, is to be preferred over mowing, but only if the flock visits each grassland patch or reserve for only a short time of less than a day, and with intervals of more than 2 weeks between visits. If mowing is preferred for other reasons, it should be done rotationally, and not before October.     

Effect of land use and climate change on the future fate of populations of an endemic species in central Europe

The identification of optimal management strategies for a given species is a major challenge of species conservation. It becomes especially challenging when the environmental conditions are expected to change in the future, and the optimal management applied today may differ from the management that is optimal under the changed conditions (e.g. due to climate change).This study evaluates prospect of a rare plant species endemic to semi-natural grasslands in central Europe, Gentianella praecox subsp. bohemica. The number of populations of this species has declined rapidly in the last 60 years; currently, a conservation action plan has been established in the Czech Republic, where most populations of this species occur. This study uses periodic matrix models to compare different management regimes under different scenarios of climate change and to identify the optimal management in each case.Without management, populations of the species are not able to survive. Flowering individuals can occur for a long time after the cessation of management, but the extinction of the population is inevitable within several decades. Without management, even very large populations (1000 flowering individuals) will go extinct in less than 50 years. Total extinction (including seed bank) will follow several years after observation of the last flowering plant. The most suitable management is mowing and disturbance (by harrowing), which is also the best method for restoration of threatened populations. Mowing is less suitable, but it is fully sufficient for large prospering populations. When managed, even small populations (10–15 flowering individuals) are able to survive. When management is applied, future climate change may have a relatively small impact on the probability of survival of the species. Climate change will, however, increase the extinction probability of very small populations.     

The impact of mowing as a management strategy for wet meadows on spider (Araneae) communities

Meadows are often managed by mowing to prevent succession and brushwood encroachment. This management practice is efficient to maintain plant diversity, but the effect on arthropod groups is less well known. We sampled spiders in the Grande Cariçaie (Switzerland), a site of nature conservation importance, in two types of wet meadows, an unmanaged site and one 2-years-old mown conditions. Numbers of individuals for the most abundant families, diversity measures, and indicator taxa were compared among vegetation and treatment types. The results indicate that the less mobile spiders and species linked to litter or dead reeds, including rare species, are reduced by mowing. Present management consists in triennial mowing of 2-4 ha non-contiguous sectors. We investigate the conflict between the need to mow the meadows to maintain them, and the negative effect of mowing on spider communities. We therefore propose a new management scheme aimed at maintain the vegetation while lessening its negative effects on spider communities by providing refuges.     

An evaluation of the status of five threatened plant species in the Pyrenees

In this paper we assess the conservation status of five of the most threatened species in the Pyrenean range (listed in the European Habitats Directive), and present updates of their distribution, reliable censuses to estimate population sizes, population growth rate, population structure, longevity, reproductive success and frequency of herbivory. Recent surveys and careful censuses revealed that the four taxa exclusively restricted to rocky habitats (Borderea chouardii, Androsace pyrenaica, Petrocoptis pseudoviscosa, Petrocoptis montsicciana) have more populations and/or individuals than previously thought. The remaining species, an orchid (Cypripedium calceolus), showed an important decline in population number. So while the rupicolous taxa might be considered naturally rare, the orchid is becoming rare. We failed to find evidence of current poor performance within populations, as recruitment was detected, population growth rate was quite stable in recent years, fruit and seed set was apparently adequate, and herbivory and predation were absent or very low. Additionally, most of the species show a long life span, which might buffer them against demographic and environmental stochasticity in absence of human disturbance.     

Long-term dynamics and population viability in one of the last populations of the endangered Spiranthes spiralis (Orchidaceae) in the Netherlands

During the last decades, most orchid species in much of Western Europe have suffered significant declines and the long-term survival of the remaining populations remains to a large extent uncertain. In particular, populations at range margins may be more prone to extinction than more central populations, as the former tend to be small and isolated, occur in ecologically marginal habitats and have a lower per-capita reproductive rate. In this study, we investigated the long-term dynamics and population viability of a population at the margin of its range of Spiranthes spiralis in the Netherlands. At present, only 2 out of 40 previously known populations persist. Individual plants were monitored for 24 years and their life span, flowering frequency and vegetative growth were determined. Individual plants showed large temporal variation in sexual and vegetative growth among years. The proportion of flowering plants varied from 0 (no plants were flowering) to 100 (all plants were flowering). Vegetative growth, on the other hand, increased when the number of individuals decreased. Dormancy was present, but occurred only in a few individuals. Using a non-structured population viability model, future prospects of this species were assessed. Calculation of extinction probabilities and estimated times to extinction using the diffusion approximation model showed that the species had a relatively high probability (79%) of surviving the next 20 years, whereas the median time to extinction was forty years. However, because 95% confidence intervals of the population growth included 1, we suggest that continued monitoring and additional genetic research are needed to assess the long-term viability of this species.     

Comparative demography of a rare species in Florida scrub and road habitats

Anthropogenic habitats can offer opportunities for expansion of rare species. The federally listed herb Hypericum cumulicola is virtually restricted to natural gaps within fire-maintained Florida scrub, but also occurs within and along sandy roads traversing scrub. To test the hypothesis that sandy roads provide suitable habitat for H. cumulicola, we compared the demographic performance of scrub and road populations at the Lake Wales Ridge State Forest and Archbold Biological Station in south-central Florida. Twice a year in February and August 1997-2006, we assessed recruitment and survival; annually in August we also measured maximum height and estimated reproductive output of tagged individuals. Scrub population dynamics were more stable than road populations. Recruitment increased with rainfall in scrub populations, but not always in road populations. Compared with scrub populations, road populations were weedier, with more variable life spans, earlier flowering, and higher fecundity. Germination rates did not differ between individuals from different habitats, but varied depending on simulated weather conditions. The weedier life history may reflect a divergent selective environment (and perhaps an evolutionary and ecological trap). Alternatively, adaptive plasticity in H. cumulicola may allow the species to maintain populations in anthropogenic habitats that can serve as refugia for fire-suppressed scrub populations.     

Variation in organic‐carbon concentration and bulk density in Flemish grassland soils

The different management regimes on grassland soils were examined to determine the possibilities for improved and/or changed land management of grasslands in Flanders (Belgium), with respect to article 3.4 of the Kyoto Protocol. Grassland soils were sampled for soil organic carbon (SOC) and for bulk density. For all grasslands under agricultural use, grazing and mowing + grazing led to higher SOC stocks compared with mowing, and grazing had higher SOC stocks compared with mowing + grazing. Overall, 15.1 ± 4.9 kg OC m^–2 for the clayey texture, 9.8 ± 3.0 kg OC m^–2 for the silty texture, and 11.8 ± 3.8 kg OC m^–2 for the sandy texture were found for grassland under agricultural use to a depth of 60 cm. For seminatural grasslands, different results were found. For both the clayey and silty texture, mowing and mowing + grazing led to higher SOC stocks compared with grazing. The clayey texture had a mean stock of 15.1 ± 6.6, the silty texture of 10.9 ± 3.0, and the sandy texture of 12.1 ± 3.9 kg OC m^–2 (0–60 cm). Lower bulk densities were found under grazed agricultural grassland compared with mown grassland but for seminatural grassland, no clear trends for the bulk density were found. The best management option for maintaining or enhancing SOC stocks in agricultural grassland soils may be permanent grazed grassland. For seminatural grassland, no clear conclusions could be made. The water status of the sampled mown fields was influencing the results for the clayey texture. Overall, the mean SOC stock was decreasing in the order clay > sand > silt. The higher mean SOC concentrations found for the sandy texture, compared to the finer silty texture, may be explained by the historical land use of these soils.     

Effects of soil properties and land management on the structure of grassland snail assemblages in SW Germany

Quantitative analyses of the snail assemblages of four a priori defined categories of grassland sites, characterized by soil moisture, land use and management, were conducted in SW Germany. Topsoil pH and calcium content were determined for each of 39 sites. The objectives were to study the effects of these habitat characteristics on snail density, species richness and quantitative structure of the land snail assemblages. With one exception, no significant relationships between soil pH (as a proxy for Ca supply) and the total numbers of individuals and species at the four site categories were observed. Snail density and species richness on groundwater soils, represented by rarely mown sedge stands and regularly mown meadows, were clearly higher than on intermediate moisture soils (as defined by the climatic water balance) of regularly mown meadows, indicating that soil moisture was the most important factor accounting for these differences. Furthermore, we observed specific conditions associated with the presence of trees in the category of meadow orchards on intermediate moisture soils. There, numbers of individuals and species were significantly higher than in the meadows on intermediate moisture soils. A posteriori cluster analysis of quantitative similarity of all snail assemblages showed that soil moisture accounts for the major differences between the site categories with respect to snail density, species richness and quantitative composition of the grassland snail assemblages. As these results are in contrast to the findings of many woodland studies, we suggest that the occurrence or abundance of certain species in the site categories are explained by land management, especially by frequent disturbances at the soil surface caused by mowing.     

Effects of management regimes and extreme climatic events on plant population viability in Eryngium alpinum

Extreme climatic events like the 2003 summer heatwave and inappropriate land management can threaten the existence of rare plants. We studied the response of Eryngium alpinum, a vulnerable species, to this extreme climatic event and different agricultural practices. A demographic study was conducted in seven field sites between 2001 and 2010. Stage-specific vital rates were used to parameterize matrix population models and perform stochastic projections to calculate population growth rates and estimate extinction probabilities. Among management regimes, spring grazing and land abandonment decreased vital rates and population growth, while autumn grazing and late mowing had positive effects on population viability. The 2003 heatwave reduced fecundity rates and survival rates. Only spring grazed sites presented considerable extinction risk. Stochastic projections showed that an increased frequency of 2003-like events may exacerbate extinction risk, but extinction probability depends mainly on land management regimes. To better conserve E. alpinum populations, we recommend conversion of presently spring grazed and abandoned sites to late mowing or autumn grazing.     

Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird

Meadow breeding birds such as the whinchat Saxicola rubetra have been declining due to increased farming intensity. In modern grassland management, the first mowing and the bird’s breeding cycle coincide, causing high nest destruction rates and low productivity of grassland bird populations. However, it is virtually unknown whether the mowing process directly affects adult survival by accidentally killing incubating females. We studied adult survival of an Alpine whinchat population during two breeding seasons using either colour-ringing or radio-tracking of 71 adults. Assessing territories, mowing phenology and nest destruction from 1988 to 2007 allowed changes in the factors associated with female mowing mortality to be estimated. Adult survival over 5-day-periods was Φ = 0.986, but during the period of mowing female survival was strongly reduced (Φ = 0.946). As a result, 80.6% of the males, but only 68.4% of the females survived the breeding season. Mowing undoubtedly killed two of 20 radio-tagged females when they were laying or incubating. In the 20-year period, an increasing proportion of nests were destroyed before the chicks hatched and this change was associated with an increased distortion of the adult sex ratio. Modelling the population growth rate showed that including the additional effect of mowing on female mortality resulted in a 1.7 times faster local population decline. These results are consistent with the hypothesis that the extinction of whinchat populations in the lowlands of central Europe was caused not only by habitat degradation and low productivity, but also by increased man-made female mortality.     

Do epiphytic orchids behave as metapopulations? Evidence from colonization, extinction rates and asynchronous population dynamics

Previous in situ studies of orchid population dynamics with conservation relevance have focused on one or a few populations in a limited area. Many species of orchids occur as hyperdispersed populations in ephemeral habitats (epiphytic, twig epiphytes, short lived or vulnerable host). In this contribution, we show that orchid populations that are patchily distributed and that exist in disturbance-prone environments may act somewhat like a metapopulation with high turnover and low correlation in population dynamics. We tested for evidence of metapopulation dynamics in the riparian orchid Lepanthes rupestris by sampling over 1000 sites (250 initially occupied, 750 initially unoccupied) in biannual surveys for 5 years. Extinction and colonization of groups of orchids on a single substrate or patch (either trees or boulders) was common and more or less consistent across different time periods, and asynchronous subpopulation dynamics were evident among the populations. From this we predict non-zero equilibrium values for site occupancy () of L. rupestris. Nevertheless, this study species differed from a typical Levins’ metapopulation system in that small populations were more likely to go extinct than large populations, and that colonization of previously occupied sites was more common than colonization of initially unoccupied sites suggesting that site quality may influence population persistence and colonization. A major difficulty applying the metapopulation approach to orchid conservation is identifying empty sites suitable for colonization. In spite of this limitation, our study highlights the necessity of following multiple orchid subpopulations (e.g., an entire orchid “metapopulation” in the broad sense) may provide a more accurate basis for predicting persistence in epiphytic orchids.     

Reduced grazing and the decline of Pulsatilla vulgaris Mill. (Ranunculaceae) in England, UK

In England Pulsatilla vulgaris is a threatened herb that declined from 130 to 33 sites between 1750 and the 1960s due to ploughing-up of calcareous grassland. We examined the subsequent fate of these populations using documentary evidence and field survey. Demographic trends were related to changes in grassland composition, structure and management and responses to increased above-ground competition (caused by reduced grazing) were simulated in a 10-year shading experiment. Since 1968 P. vulgaris has been lost from 16 sites and gradually declined on four others. However, the total population size increased by 258% due to the reintroduction of winter grazing on three sites. This produced a significantly shorter, more herb-rich sward, with a lower cover of Bromopsis erecta (c. 10%), than on sites where populations remained stable or declined. Experimental shading had a significant negative effect on plant survival and flowering performance. These results confirmed that reduced grazing is now one of the major threats to species dependent on short swards, especially on isolated sites where livestock farming is no longer economically viable. That many of these declines took place on nature reserves highlights the difficulties of managing isolated grasslands, and the urgent need to re-instate grazing on reserves supporting populations of threatened species in otherwise intensively managed lowland landscapes.