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.     

Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment

Calcareous grasslands, most of which are man made and therefore depend on some kind of human interference for their maintenance, are among the most species-rich communities on Earth at a small scale. For many centuries, most of these grasslands have been used as extensive pasture. However, after 1900, and particularly from 1940 onwards, livestock grazing has declined throughout Europe leading to the abandonment of low intensity grasslands over large areas. To conserve the remaining grasslands or to restore recently abandoned grasslands, better insights about the effects of grassland management on above and belowground species diversity are needed. Here, we describe the results of an 11-year experiment to investigate the role of grassland management (grazing, mowing and abandonment) in determining species composition and diversity both in the aboveground vegetation and the seed bank of a calcareous grassland in Belgium. Species diversity declined by about 60% 11 years after abandonment, from 29 species m⁻² to as few as 12 species m⁻². Plots that were grazed remained constant in species richness, whereas mown sites lost about 20% of their original species. Abandoned plots were largely dominated by a few grass species, in particular Festuca rubra. Concomitant with changes in the aboveground vegetation, both the number of species found in the seed bank and seed density (number of seeds m⁻²) had changed significantly 11 years after abandonment. Species diversity and seed density were significantly lower in abandoned plots than in grazed or mown plots. We conclude that abandonment of calcareous grasslands may lead to rapid decline of plant species diversity both in the aboveground vegetation and in the seed bank. As a result, seed banks probably have a limited role to play in the restoration of recently abandoned grasslands.     

Does nectar reward affect rarity and extinction probabilities of orchid species? An assessment using historical records from Belgium and the Netherlands

The long-term persistence of plant populations may partly depend on pollination processes and seed production. The exact role of pollination in determining plant population viability still remains largely unclear. Orchid species have often been shown to be pollinator limited. Especially, non-rewarding species are characterized by infrequent pollinator visits and low seed set. Therefore, it can be hypothesized that, if overall population fitness of orchid species is influenced by seed production, non-rewarding species should be more prone to local extinction than rewarding species. To study the importance of nectar reward on orchid persistence, we collected historical records about the distribution of 32 orchid species in Flanders (Belgium) and 37 orchid species in the Netherlands. For both regions, present distribution patterns were compared with historical data. Between 1930 (Flanders) or 1950 (the Netherlands) and 2000, 26 (81%) species showed decreases in distribution range in Flanders, eight of which had gone extinct, whereas 29 (78%) species showed declines in distribution area in the Netherlands, five of which went extinct. Contrary to previously reported results, orchid distribution patterns were not related to nectar reward. There was also no significant relationship between nectar reward and extinction for both regions. Orchids typically occurring in wet grasslands and heathlands suffered greater losses than orchids typically confined to forest habitats or calcareous grasslands. We conclude that the production of nectar does not represent a safeguard for local extinction of orchid species. Habitat loss and other deterministic threats associated with habitat fragmentation and deterioration are more important determinants of orchid persistence.     

Relative effects of management and environmental conditions on performance and survival of populations of a terrestrial orchid, Dactylorhiza majalis

For population viability analysis of endangered orchid populations, it is crucial to disentangle the effects of weather and management from intrinsic orchid dynamics. When doing this, typically several months’ average temperatures and/or sums of precipitations are compared with some characteristics of plant performance. Here we tested, whether short averaging intervals (1-2 weeks) are more closely correlated with orchid performance. We used 5 years of data from five Dactylorhiza majalis populations, and have shown that the improvement of prediction by shortening the interval over which the temperatures are averaged or precipitation summed, even if detectable, is only weak and not significant. This, however, may be due to low weather variability during the study. Regarding the second aspect, the effect of management (presence or absence of mowing), we have found that leaf area of D. majalis at the regularly mown site was larger than that at the sites which were mown only once in 2 years, but we did not detect a significant effect of the absence of mowing on the incidence of flowering. Mowing can affect orchid performance in two ways: by reduced shading of orchids and by reducing competition with other species. Therefore, we have determined the co-occurring species associated with presence or absence of mowing and found that shading significantly affected the length of the flower stalk, the ratio of leaf width to leaf length at the end of the season, but did not affect seed weight and probability of flowering the next year. We conclude that the most appropriate management for D. majalis is mowing at least once a year, ideally at the end of June/beginning of July, after its fruiting.     

Influence of habitat fragmentation on the genetic structure of Polyommatus coridon (Lepidoptera: Lycaenidae): implications for conservation

Conservation biologists pay increasing attention to the interdependence between habitat structure and genetic patterns of species and to the importance of conserving genetic diversity. A suitable model species for such a study is Polyommatus coridon, a butterfly of dry calcareous grassland. We studied the population genetic structure of 22 western German populations of this species (874 individuals). We observed no congruence between the observed genetic structure and the geographic arrangement of the studied populations. Genetic diversity was slightly higher in large populations than in small populations. Differentiation among populations was weak or even missing. Populations of an area with high habitat density showed no significant differentiation, whereas areas with low habitat densities showed significant genetic differentiation among populations. At present, hierarchical variance analysis revealed no differentiation between four distinguished limestone areas. The combination of all analytical results allow the definition of management units for P. coridon. Conservation measures are proposed from which a large number of rarer species occurring together with P. coridon will benefit.     

The requirements of the rare moss, Hamatocaulis vernicosus (Calliergonaceae, Musci), in the Czech Republic in relation to vegetation, water chemistry and management

Hamatocaulis vernicosus, a rare moss, has been investigated in detail for its habitat preferences, ecology and population dynamics in the Czech Republic. At all its known sites plant species composition was described and relationships with environmental factors investigated (water table, pH, water conductivity). Experiments that included mowing and gap cutting were investigated at three sites over two years.Hamatocaulis vernicosus had the highest cover at neutral pH (6.7-7.2) and conductivity between 100 and 250 μS/cm, although most localities had lower values. It was influenced positively by mowing only at a site with a high vascular plants cover, and gap cutting was only beneficial at sites with a low water table. The growth and vitality of Hamatocaulis may, therefore, be supported by suitable management especially in drier habitats.     

Effects of fragmentation on pollinator assemblage, pollen limitation and seed production of Mediterranean myrtle (Myrtus communis)

Pollinator assemblages may shift as a consequence of the destruction and fragmentation of natural habitats. The scarcity of mates and pollinators can lead plant populations to suffer from pollen limitation and a decrease in reproductive performance within fragmented areas. We studied the shift in pollinator assemblages along with pollen limitation and seed production patterns in the Mediterranean shrub Myrtus communis. Our study included six populations contrasting in patch and population size (Large vs. Small) within a fragmented landscape characterized by ∼1% of potential forest coverage. The breeding system in Myrtus communis was self-compatible, but compared with natural pollination, fruit set increased with pollen addition (quantity limited), and seed set (brood size) increased with outcross pollen addition (quality limited). While the pollinator assemblage in Large patches was taxonomically diverse, it was almost monopolized by honeybees in Small patches, where visitation rates were highest and wild bee species were almost absent. In general, Small populations were less pollen limited for fruit set than Large populations, particularly those that received the highest rates of honeybee visits. However, despite differences in fragmentation and pollinators between Large and Small populations, seed production patterns (brood size and seed mass) were rather similar among them, in agreement with similar pollen limitation levels found for brood size. A higher susceptibility of native pollinators to the presence of honeybee hives was found in Small patches, suggesting that the pollinator assemblage may be severely altered when fragmentation occurs in combination with beekeeping. We discuss its implications and effects on plant reproduction in fragmented areas.     

Low impact of present and historical landscape configuration on the genetics of fragmented Anthyllis vulneraria populations

Decreasing habitat fragment area and increasing isolation may cause loss of plant population genetic diversity and increased genetic differentiation between populations. We studied the relation between the historical and the present landscape configuration (i.e., patch area and patch connectivity), and the present management of calcareous grassland fragments on the one hand, and the within and between population genetic structure of 18 Anthyllis vulneraria populations on the other hand. Despite the long-time fragmentation history and the mainly selfing breeding system of the species, we detected very low genetic differentiation (Φ_st = 0.056) among habitat fragments and no significant isolation-by-distance relation. Average within fragment genetic diversity measured as molecular variance and expected heterozygosity, were relatively high (16.46 and 0.28, respectively), and weakly positively correlated with the current fragment area, most likely because larger fragments contained larger populations. We found no effects of the historical landscape configuration on the genetic diversity of the populations. Our data suggest that the consequences of habitat fragmentation for genetic differentiation and genetic diversity of A. vulneraria are relatively minor which is very likely due to the historical high levels of seed exchange among fragments through grazing and roaming livestock. This study provides indirect evidence that nature management by grazing not only positively affects habitat quality but that it might also mitigate the genetic consequences of habitat fragmentation. From the conservation point of view, this study illustrates the importance of grazing and of the regular transport of livestock between fragments to prevent the long-term effects of fragmentation on the genetic diversity of the populations studied.     

Pollination of the invasive exotic shrub Lupinus arboreus (Fabaceae) by introduced bees in Tasmania

Exotic plant invasions threaten ecological communities world-wide. Some species are limited by a lack of suitable pollinators, but the introduction of exotic pollinators can facilitate rapid spread. In Tasmania, where many non-native plants are naturalised, exotic honeybees (Apis mellifera) and bumblebees (Bombus terrestris) have become established. We determined how these species affect the pollination of Lupinus arboreus, an invasive, nitrogen-fixing shrub, which is rarely visited by native pollinators. The proportion of flowers setting seed and the number of ovules fertilised per flower were positively related to the visitation rates of both exotic bee species. There was no effect of bee visitation rates on the proportion of seeds aborted prior to maturity, possibly due to post-fertilisation environmental constraints. We conclude that the spread of B. terrestris may not alter the fecundity of L. arboreus because of the pollination service provided by A. mellifera, and discuss potential interactions between these two bee species.     

Conservation management of calcareous grasslands. Changes in plant species composition and response of functional traits during 25 years

Five management treatments have been applied to a calcareous grassland since 1974 to study their efficiency in maintaining high diversity of grassland species. The treatments were: continuing sheep grazing, mowing, mulching, burning and leaving fallow (unmanaged). After 25 years of management, the treatments mowing and mulching resulted in a species composition similar to that from grazing by sheep. Thus, mowing and mulching can be useful management alternatives to grazing. In contrast, both burning and leaving fallow considerably changed the species composition. The changes were similar in both treatments and neither can be recommended for management purposes. We also evaluated eight traits with assumed sensitivity to the treatments (life form, life cycle, growth form, runners, lateral spread, fecundity, seed mass, germination season). In 1999, all traits except life cycle were found to respond to the different treatments. Trait response is discussed in detail.     

Breeding system and inbreeding depression in the clonal plant species Knautia arvensis (Dipsacaceae): implications for survival in abandoned grassland

Larger clonal sizes observed in abandoned than in managed grassland could result in increased geitonogamy in Knautia arvensis. The species was investigated for self-compatibility, seed set without pollinators, and inbreeding depression in a greenhouse experiment. Seed set was equivalent in hand-selfed and hand-outcrossed plants (c. 53%), indicating full self-compatibility. Seed set was low (2.8%) in caged, unmanipulated flower heads. There was no significant difference in seed weight, germination, and survival between selfed and outcrossed progeny. Juvenile biomass was significantly lower in selfed than in outcrossed progeny (on average 45% lower). Cumulative fitness calculations for germination, survival, and juvenile biomass showed that the mean inbreeding depression was 58% in offspring produced by selfing. The results suggest that in K. arvensis, which is protected against self-pollination within flower heads through protandry, the likely mechanism for selfing is via geitonogamy among flower heads. Geitonogamy among ramets within the larger clones observed in abandoned grassland could reduce offspring viability in K. arvensis since the fitness of selfed offspring was low. Inbreeding depression could therefore add to the recruitment problems experienced by plants in abandoned grassland, both in K. arvensis and probably in other clonal grassland species, depending on their growth structure and breeding system.     

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.     

Effect of the presence of nectar feeders on the breeding success of Salvia mexicana and Salvia fulgens in a suburban park near México City

The use of nectar feeders has grown a lot as a result of people wanting to attract hummingbirds to their gardens. This has caused an increment in the density and diversity of these birds in urban and suburban habitats. The feeders not only represent a bonanza resource for hummingbirds but they can also result in reduced visitation to native plants. We experimentally assessed the effect of placing nectar feeders in the vicinity of two native plant species in a suburban area of Mexico City. The plants with feeders showed a reduced visitation rate and in one case (Salvia fulgens) produced fewer seeds than the controls. Salvia mexicana was not only visited by hummingbirds but it was also by bees that were not attracted to the feeders, probably resulting in the lack of effect on this plant. Native plants in urban and suburban habitats can be threatened by reduced visitation rates and reduced seed sets resulting from the use of feeders.     

Ecological and genetic effects on demographic processes: pollination, clonality and seed production in Dithyrea maritima

The relative importance of ecological and genetic factors in regulating the demographic processes of small populations is unknown for most rare plants. Dithyrea maritima (beach spectacle pod; Brassicaceae) is a rare, rhizomatously spreading, and self-incompatible herbaceous perennial endemic to coastal sand dunes of southern California. Low seed production in this species may be caused by insufficient pollinator visits, which may be exacerbated by competition for pollinator visits from non-native plants that have invaded the coastal dunes. I tested this ‘ecological’ hypothesis by removing flowers of Cakile maritima and Carpobrotus spp. from three experimental plots on San Nicolas Island. A second possible limitation on seed set is low diversity of genets within subpopulations, which could lead to scarcity of compatible mates. I tested this ‘genetic’ hypothesis by assigning ramets to clones using isozymes, and by relating clonal diversity within populations to seed set. Removal of non-native plants had no detectable effect on pollination service to D. maritima or on seed set, and seed set was unrelated to natural variation in pollination service among populations within a single year. In contrast, clonal diversity within a 10 m² patch was a strong predictor of seed set. Hence a genetic factor explains spatial variation in seed set on San Nicolas Island. On larger spatial and temporal scales, however, pollination service also affects patterns of seed set: a shift in pollination from flies in 1998 to bees in 1999 probably caused a corresponding increase in seed set across San Nicolas Island, while extremely low seed set at another site (San Miguel Island) is probably caused by a virtual absence of pollinators there.     

Optimizing grassland management for flower-visiting insects in roadside verges

The decline of flower-visiting insects is a threat to ecological processes and to the services these insects provide. Roadside verges in the Netherlands span approximately 80,000 km and are often covered with semi-natural grasslands. As such, they also provide a suitable habitat for many insects, but this has received little attention so far. We investigated the effects of different management treatments on flower-visiting insects. We studied flower visitation in a 3 years old experimental set-up with five mowing treatments each replicated five times. Management types were: no management and mowing once or twice per year with and without the removal of hay, representing common forms of management and neglect. During an entire growing season, both flowers (number of species and inflorescences) as well as insects (total abundance and actual flower visits) were investigated. Mowing twice per year with removal of hay showed highest values for all measured variables and this effect persisted throughout the growing season. The early summer cut proved to be very important for insect feeding opportunities, due to the re-flowering of plants later in the growing season. Flower abundance showed high correlations with both plant species richness and the number of insect visits. Although overall, mowing twice a year with hay removal was the most beneficial treatment for flower-visiting insects, these plots were entirely devoid of flowers for some period right after mowing, indicating that a rotational scheme might further promote insect diversity and abundance.     

The bryophyte layer in a calcareous grassland after a decade of contrasting mowing regimes

Analysis of variance and redundancy analysis were used to study the effects of mowing regime and herb canopy structure on the bryophyte layer of a calcareous grassland in southern Belgium. Species richness in mown and control plots did not significantly differ. In control plots, a number of grassland species were able to survive within a grassy-shrubby fringe 15 years after shrub cutting. This suggests that there might be a time-lag of at least 20 years before cessation of mowing results in a significant decrease in species number per m². Mown plots were mostly characterized by the almost complete cover of large pleurocarps. Mowing resulted in almost continuous herb and moss layers that were unfavourable for a number of annual bryophytes, many of which are of high conservation interest. It is suggested that grazing, which increases the number of species, especially the number of pioneers, is likely to be more beneficial to bryophytes.     

Herbivore and pollinator responses to grassland management intensity along experimental changes in plant species richness

Agricultural intensification is a major driver of global environmental change. Disentangling the relative impact of losses in plant species richness and intensified management on higher trophic level organisms is important for conservation recommendations.We established different management regimes along an experimental gradient of plant species richness within “The Jena-Experiment” in Germany and quantified herbivory as well as grasshoppers and pollinators. Herbivory, grasshopper density and species richness and frequency of flower visiting pollinators were recorded two times in each of four subplots of altogether 80 plots differing in plant species richness. Each of the four subplots was subject to four different levels of fertilizer application and mowing to simulate very low, low, high and very high land use intensity.Fertilization and mowing significantly affected plant–herbivore interactions but plant species richness had no discernible effect. Grasshoppers were most abundant at high intensity subplots (3.1 individuals per m²) and least abundant on very low intensity subplots (1.3 individuals per m²). Leaf damage caused by herbivores was highest in even the very high intensity subplots (3.7%) with four mowing events per year and high amounts of fertilizer application and lowest on subplots with a low management intensity (2.4%) comprising two mowing events per year but no fertilizer application. In contrast, pollinators benefited most from lower management intensities, with only one or two mowing events and no fertilizer addition. In addition, higher numbers of flowering plant species and increased blossom cover was associated with enhanced pollinator species richness and flower visitation.Our results indicate that even in grasslands with high plant species richness, mowing and fertilization are more important drivers of herbivory and flower visitation by pollinators. Management with no more than two cuts per year and without fertilizer application in our grasslands balanced the ecosystem functions of increased pollination and decreased herbivory.     

Conservation of the red-winged grasshopper, Oedipoda germanica (Latr.): the influence of reproductive behaviour

Reproductive isolation can function as a mechanism to maintain locally adapted gene complexes while decreasing the heterozygosity in distinct populations. As a result, reproductive behaviour should be considered a fundamental factor influencing reproductive isolation. This is of interest to conservation biology when one desires to regulate gene flow between two populations either by creating opportunities for increased dispersal, by relocation of individuals, or by re-introduction of a species to its natural habitat. Reproductive behaviour can also influence the effective population size and the actual population size through the Allee effect. We investigated the reproductive behaviour of individuals from two isolated populations of the red winged grasshopper, Oedipoda germanica, an endangered species in Central Europe. We detail several methods to show how several aspects of the reproductive behaviour of this species interact with the conservation of this species. Foreign males were not disfavoured in mate choice and male body size was also unimportant in mating success. Heterogamic matings were as productive as homogamic matings in terms of total number of eggs per female, egg hatching rate, or nymph survival. Therefore, we suggest that cross-matings of individuals from different populations do not positively influence population size by heterosis effects nor act they negatively in the form of outbreeding depression. We found that female O. germanica were able to store viable sperm for extended periods but egg pods showed a decrease in hatching rate when these females were deprived of additional mating opportunities. Multiple mated females laid more eggs than once-mated females. Hence, females are capable of founding new populations even after only one mating but their reproductive output may be reduced. Present-day gene flow between populations of O. germanica probably does not occur. Consequently, there is an urgent need to pursue habitat management and release programmes that maintain current population sizes of this species. With regard to the reproductive behaviour of the red-winged grasshopper, we suggest that relocation programmes release males and female together and at an early adult stage. Also, because female reproductive output increases with mating activity, the initial release should involve excess females. Because males have a shorter lifespan than females a subsequent release of males at a later time may enhance the overall success of local conservation efforts by resulting in more offspring per female.     

Effects of excluding grazing animals from grassland on sugar limestone in Teesdale,England

The effects of excluding sheep and rabbits for 12 years from grassland on sugar-limestone on Cronkley Fell, Teesdale in the north Pennines, are described. Frequency measurements from permanent quadrats inside the enclosure and from control quadrats outside, show that dwarf shrubs, particularly the nationally rare species Dryas octopetala and Helianthemum canum, benefit from excluding grazing. The grassland structure has slowly changed to become more open, mainly through the death of Festuca ovina and reduction in bryophytes and fruticose lichens, but only one species, Gentianella amarella, has been lost. Exclosure of grazing animals is therefore shown to be a succesful management procedure for this community where plant growth is restricted by severe climate and limiting soil-nutrient factors.     

Reproductive biology and ecology of selected rare and endangered Oxalis L. (Oxalidaceae) plant species

About twenty-five percent of all southern African Oxalis species are rare/endangered and highly localized, making them especially vulnerable to extinction through inbreeding, low genetic variation, disrupted biological interactions and stochastic events; all consequences of small population sizes. Moreover, Oxalis displays tristyly, which is a rare and specialized sexual system that includes a strong self-incompatibility component between three floral morphs to promote out-crossing within populations. As tristyly requires the availability of plants with different floral morphs as well as effective pollinators for seed production, this breeding system can affect small populations when fully expressed. Factors that may have an effect on rarity in Oxalis were investigated by focusing on the expression of tristyly, levels of natural seed production, clonality and the ecology of eight rare/highly localized Oxalis species. Field experiments revealed that the reproductive success of some Oxalis species may be hampered by tristyly, resulting in extremely low levels of natural seed production. Other species display a more relaxed expression of self-incompatibility, which in combination with the possibility of cross-pollinations provides reproductive assurance regardless of population structure and pollinator availability. Others are rare and endangered, but appear not to be negatively affected by the tristylous breeding system. Most species are limited by their highly specific habitat requirements and are particularly vulnerable to variation in rainfall patterns.