Comparison of reproductive success in two orchids: the nectarless Dactylorhiza majalis s.s. and the nectar-producing Gymnadenia conopsea s.l.

Differences in reproductive biology between two orchid species were examined: Eight Danish populations of the nectarless Dactylorhiza majalis ssp. majalis and six Swedish populations of the nectar-producing Gymnadenia conopsea ssp. densiflora . Population size varied from 24 to 1700 flowering individuals in D. majalis and from 100 to 1200 in G. conopsea. Dactylorhiza majalis had only half as many flowers per spike as G. conopsea . Fruit-set of D. majalis ranged from 16 to 39%, much lower than the fruit-set of G. conopsea (78–91%). Thus in the species with a few flowers and no nectar a lower fruit-set was observed than in the species with many flowers and presence of nectar. The low fruit-set of D. majalis was pollen-limited and fruit-set was positively correlated with pollinia removal and the latter was negatively correlated with population size. In both species, fruit-set and population size were uncorrelated.     

Effects of population size and forest management on genetic diversity and structure of the tuberous orchid Orchis mascula

Due to societal changes and altered demands for firewood, the traditional forest management of coppicing has been largely abandoned. As a result, many forest herbs that are specifically adapted to regular opening of the canopy, have suffered significant declines in abundance, and the remaining populations of these species often tend to be small and isolated. Reduced population sizes and pronounced spatial isolation may cause loss of within-population genetic diversity and increased between-population differentiation through random genetic drift and inbreeding. In this study, we investigated genetic diversity and genetic structure of 15 populations of the food-deceptive orchid Orchis mascula using AFLP markers. Within-population genetic diversity significantly increased with increasing population size, indicating genetic impoverishment in small populations. Genetic differentiation, on the other hand, was rather low (Φ_ST = 0.083) and there was no significant relationship between genetic and geographic distances, suggesting substantial gene flow within the study area. However, strong differences in levels of within-population diversity and among-population differentiation were found for populations located in forests that have been regularly coppiced and populations found in forests that were neglected for more than 50 years and that were totally overgrown by shrubs. Our data thus indicate that a lack of coppicing leads to decreased genetic diversity and increased differentiation in this orchid species, most likely as a result of genetic drift following demographic bottlenecks. From a conservation point of view, this study combined with previous results on the demography of O. mascula in relation to forest management illustrates the importance of coppicing in maintaining viable populations of forest herbs in the long-term.     

Patterns of genetic variation in Gymnadenia conopsea, the fragrant orchid

Gymnadenia conopsea (L.) R. Br., or the fragrant orchid, is one of many plant species negatively influenced by new practices in agriculture and forestry during the last decades. This study describes the level of microsatellite variation within and among 10 Swedish populations of this species. It was not possible to detect strong effects of small population size or fragmentation. In general, the species had high genetic variation within and low genetic divergence among populations, although the correlation between population size and number of alleles was close to significance at the 95% level. Also, a significant isolation by distance effect was observed, indicating the presence of modest restrictions in gene dispersal between the investigated populations.     

不同培养基对兰科药用植物手参原球茎共生真菌的分离效果

兰科植物手参Gymnadenia conopsea作为国家二级重点保护野生植物具有重要的药用价值。目前手参还未实现人工栽培,但其种子的真菌共生萌发已获成功。为明确除促萌发真菌外,还有哪些土著真菌参与了手参种子的萌发过程,本研究在自然条件下采用促萌发真菌伴播手参种子,获得了种子萌发形成的原球茎,进而对比了6种常见的培养基PDA (马铃薯葡萄糖琼脂培养基)、MMN (改良Melin-Norkrans培养基)、FIM (真菌分离培养基)、MEA (麦芽浸膏琼脂培养基)、CAM (胡萝卜葡萄糖琼脂培养基)和CMA (玉米粉琼脂培养基)对手参原球茎共生真菌分离效果的影响。共从6种培养基上分离获得了75个菌株,其中MMN、CAM、PDA、FIM、MEA、CMA培养基依次分离得到20株、16株、15株、11株、8株、5株菌。此外,真菌的多样性分析结果表明,MMN培养基的Chao 1、Shannon-Wiener和Simpson多样性指数最高,CAM和PDA培养基次之,CMA培养基最低。综上所述,真菌分离效果最好的是MMN培养基,其次是CAM和PDA培养基,而FIM和MEA培养基对真菌的分离效果影响不大,CMA培养基的分离效果最差。本研究结果可为其他兰科植物原球茎共生真菌的分离提供借鉴,所获得的菌株也有望进一步应用于功能菌剂的研发。     

Genetic differentiation and habitat preference of flowering-time variants within Gymnadenia conopsea

Using fast-evolving microsatellites, more slowly evolving ITS markers and performing habitat analyses, we demonstrated a drastic genetic divergence and significant habitat differentiation between early- and late-flowering variants of plants morphologically belonging to Gymnadenia conopsea ssp conopsea. The two phenological variants can either be found in separate or in mixed populations. Information from microsatellite markers and ITS sequences indicated the occurrence of an early historical split between the two flowering-time variants, a split that has been maintained until the present time even within sympatric populations. Early-flowering variants were also far more genetically diverse, had more alleles per microsatellite locus and a wider habitat amplitude than late-flowering variants. As a comparison, we included G. odoratissima in the sequencing study. We found G. odoratissima to be most closely related to the early-flowering type. This indicates a more ancient divergence event between the two flowering-time variants within G. conopsea ssp conopsea than between the two different species G. odoratissima and the early-flowering variant of G. conopsea. Possible explanations to the results arrived at and possible mechanisms maintaining the genetic separation are discussed.     

Phenotypic selection on nectar amino acid composition in the Lepidoptera pollinated orchid species Gymnadenia conopsea

Plant–pollinator interactions are potential drivers of evolution in floral traits. Because nectar chemical composition is known to mediate both plant–pollinator interactions and plant reproductive success, it can be expected that chemical composition of nectar is subjected to strong pollinator‐mediated selective forces. However, the extent of natural selection on different nectar components has not been studied so far. Using the Lepidoptera pollinated fragrant orchid Gymnadenia conopsea as a model species, we used high‐performance anion‐exchange chromatography (HPAEC) to characterize the sugar and amino acid composition of floral nectar in three calcareous grassland populations of G. conopsea. We then measured phenotypic selection on nectar composition and on other plant and floral traits through applying both linear regression and structural equation modelling. We demonstrate phenotypic selection on plant height, inflorescence height and on specific nectar amino acids, whereas spur length, total sugar and amino acid concentration were not direct targets of selection. Chemical nectar composition is thus indeed under selective pressure but nectar amino acids are much more important to fitness of G. conopsea, as compared to nectar sugars. Furthermore, as we found no evidence of selection on the total amino acid concentration, it is unlikely that amino acids increase pollinator attraction because they are a pollinator nitrogen source. To further unravel the evolutionary ecology of floral nectar, behavioural experiments with pollinators exposed to different nectar components and studies experimentally identifying the selective agents are recommended.     

Among-population variation in costs of reproduction in the long-lived orchid Gymnadenia conopsea: an experimental study

A cost of reproduction in terms of reduced future performance underlies all life-history models, yet costs have been difficult to detect in short-term experiments with long-lived plants. The likelihood of detecting costs should depend on the range of variation in reproductive effort that can be induced, and also on the shape of the cost function across this range, which should be affected by resource availability. Here, we experimentally examined the effects of both reduced and increased fruit production in two populations of the long-lived orchid Gymnadenia conopsea located at sites that differ in length of the growing season. Plants that were prevented from fruiting produced more flowers in the population with a longer growing season, had higher survival in the other population, and grew larger compared to control plants in both populations. Fruit production was pollen-limited in both populations, and increased reproductive investment after supplemental hand-pollination was associated with reduced fecundity the following year. The results demonstrate that the shape of the cost function varies among fitness components, and that costs can be differentially expressed in different populations. They are consistent with the hypothesis that differences in temporal overlap between allocation to reproduction and other functions will induce among-population variation in reproductive costs.     

Genetic diversity in the rare orchid Gymnadenia odoratissima and a comparison with the more common congener, G. conopsea

Patterns and levels of genetic diversity mayhave significant influence on the long termpersistence of local populations and revealingsuch information is important in protectingrare species. In this study we investigated thegenetic pattern in five microsatellite lociwithin five Swedish populations of the rareorchid species Gymnadenia odoratissima. Thegeographic distribution of G. odoratissima isrestricted to Europe and in Scandinavia it isonly found in three provinces in southernSweden; Östergötland,Västergötland and on the island ofGotland.Compared with the more widespread congener G.conopsea our results indicate lower levels ofgenetic variation within and higher degrees ofgenetic differentiation among populations ofG. odoratissima (H_EL = 0.6–0.8 in G. conopseaand 0.3–0.7 in G. odoratissima; F_ST over allpopulations = 0.06 in G. conopsea and 0.19 inG. odoratissima). Also, we found a cleardistinction among mainland and islandpopulations of G. odoratissima wherepopulations on the island of Gotland seem toexhibit higher levels of gene flow andintragenetic variation, probably as a result ofa larger number of existing populations.Future conservation of this species shouldfocus on facilitation on colonisation events,especially on the mainland, and preservation ofthe genetically more variable Gotlandpopulations.     

Remarkable coexistence of multiple cytotypes of the Gymnadenia conopsea aggregate (the fragrant orchid): evidence from flow cytometry

Background and Aims

One of the prerequisites for polyploid research in natural systems is knowledge of the geographical distribution of cytotypes. Here inter- and intrapopulational ploidy diversity was examined in the Gymnadenia conopsea aggregate in central Europe and potential explanations and evolutionary consequences of the observed spatial patterns investigated.

Methods

DAPI flow cytometry supplemented by confirmatory chromosome counts was used to determine ploidy in 3581 samples of the G. conopsea aggregate from 43 populations. The fine-scale spatial pattern of cytotype distribution (intra- and interploidy associations) was analysed with univariate and bivariate K-functions.

Key Results

Gymnadenia tissues undergo a progressively partial endoreplication, which accounts for about 60 % and 75 % of the total genome in G. conopsea and G. densiflora, respectively. Flow cytometric profiles are therefore species-specific and can be used as a marker for rapid and reliable species recognition. Two majority (4x, 8x) and three minority (6x, 10x, 12x) cytotypes were found, often in mixed-ploidy populations (harbouring up to all five different ploidy levels). The scarcity of the minority cytotypes (about 2·7 %) suggests the existence of strong pre- or postzygotic mating barriers. Spatial structure was observed in plots of populations with the highest cytotype variation, including clumping of individuals of the same ploidy and negative association between tetra- and octoploids.

Conclusions

The remarkable ploidy coexistence in the G. conopsea aggregate has reshaped our perception of intrapopulational ploidy diversity under natural conditions. This system offers unique opportunities for studying processes governing the formation and establishment of polyploids and assessing the evolutionary significance of the various pre- and postzygotic mating barriers that maintain this ploidy mixture.     

Floral display and habitat fragmentation: Effects on the reproductive success of the threatened mass‐flowering Conospermum undulatum (Proteaceae)

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Effects of individual and population parameters on reproductive success in three sexually deceptive orchid species

Reproductive success (RS) in orchids in general, and in non-rewarding species specifically, is extremely low. RS is pollinator and pollination limited in food deceptive orchids, but this has rarely been studied in sexually deceptive orchid species. Here, we tested the effects of several individual (plant height, inflorescence size, nearest neighbour distance and flower position) and population (patch geometry, population density and size) parameters on RS in three sexually deceptive Ophrys (Orchidaceae) species. Inter-specific differences were observed in RS of flowers situated in the upper versus the lower part of the inflorescence, likely due to species-specific pollinator behaviour. For all three species examined, RS increased with increasing plant height, inflorescence size and nearest neighbour distance. RS generally increased with decreasing population density and increasing patch elongation. Given these results, we postulate that pollinator availability, rather than pollinator learning, is the most limiting factor in successful reproduction for sexually deceptive orchids. Our results also suggest that olfactory 'display' ( i.e. versus optical display), in terms of inflorescence size (and co-varying plant height), plays a key role in individual RS of sexually deceptive orchids. In this regard, several hypotheses are suggested and discussed.     

Pollen fates and the limits on male reproductive success in an orchid population

Male reproductive success in higher plants depends largely on the fates of pollen, but current methodologies have given only partial insights into this important aspect of plant mating. We present a detailed analysis of the proportions and absolute amounts of stained pollen involved in six key fates for the hawkmoth-pollinated African orchid Disa cooperi . Despite being packaged into pollinaria, high proportions (> 0.95) of the pollen removed from anthers were lost during transport by hawkmoths in both years. The proportion of pollen lost correlated positively with the number of pollinaria removed from a plant, so that pollen export did not vary with pollen removal. Most pollen was dispersed to neighbouring plants, with rare long-distance dispersal up to 65 m. Of the pollen that reached stigmas during both years, roughly equal amounts were involved in facilitated self-pollination vs. cross-pollination, but the relative proportions of these fates differed between years. Contrary to expectation, we found that self-pollination between flowers did not increase with the number of open flowers, even though moths probed significantly more flowers on larger plants. However, during both years the fraction of removed pollen exported to other plants declined significantly with increasing self-pollination on the source plant, indicating that once self-pollination occurred it reduced (discounted) subsequent pollen export opportunities. The packaging of pollen into pollinaria in orchids appears to increase overall transfer efficiency by at least an order of magnitude relative to plants with granular pollen. Nevertheless, considerable uncertainties remain in the male reproductive success of individual orchids. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 175–190.     

生境破碎化对生物多样性的影响

生境破碎化对生物多样性和生态系统功能的影响是当前国内外生态学家研究的热点问题之一。生境破碎化导致原生境的总面积减小,产生隔离的异质种群,从而影响个体行为特性、种群间基因交换、物种间相互作用及生态过程。生境破碎化的过程引起栖息地内部食物、繁殖场所、局部小气候、边缘效应等生物和非生物条件的变化,从而影响植物种群的大小和灭绝速率、扩散和迁入、遗传和变异以及存活力等,影响动物种群的异质种群动态、适宜生境比例、灭绝阈值、种间关系等。随着景观生态学与农业科学的融合,探索利用景观布局控制害虫发生将是人类利用生境破碎化为人类服务的一条新途径。     

Effects of local and landscape-scale habitat variables on abundance and reproductive success of wetland birds

Both local and landscape-scale habitat variables influence the abundance of wetland breeding birds. Few studies, however, simultaneously assess the effects of habitat variables at multiple spatial scales or consider effects on reproductive success. Therefore, we examined the effects of wetland and landscape-scale habitat variables on the abundance of nine breeding bird species and the effects of nest, wetland, or landscape-scale habitat variables on the nest success, clutch size, or number of fledglings of four species at 15 cattail (Typha sp.)-dominated wetlands in an agricultural region around Peterborough, Ontario, Canada. The abundance of Least Bittern (Ixobrychus exilis), Common Moorhen (Gallinula chloropus), and Marsh Wren (Cistothorus palustris) increased as wetland water depth increased; the abundance of Common Moorhen and Marsh Wren increased as wetland size increased; and the abundance of Marsh Wren increased as the amount of wetland in the surrounding landscape increased. Red-winged Blackbird (Agelaius phoeniceus) nest success decreased as nest cover increased. Clutch sizes were uninfluenced by the habitat variables that we considered. The number of Red-winged Blackbird fledglings per successful nest increased as wetland size increased and as the amount of wetland in the surrounding landscape increased. We speculate that food limitation in small wetlands may be responsible for the pattern in Red-winged Blackbird fledging success. The abundance and nest success of Virginia Rail (Rallus limicola) and Sora (Porzana carolina) were uninfluenced by the habitat variables we considered. Future research should consider mate attraction and productivity in relation to local and landscape-scale habitat variables for these and other secretive species. Our study suggests that wetland conservation will be most effective if it considers habitat variables at multiple spatial scales.     

Ecological interactions of the sexually deceptive orchid Orchis galilaea

Plant species dependent on highly specific interactions with pollinators are vulnerable to environmental change. Conservation strategies therefore require a detailed understanding of pollination ecology. This two-year study examined the interactions between the sexually deceptive orchid, Orchis galilaea, and its pollinator Lasioglossum marginatum. Relationships were investigated across three different habitats known to support O. galilaea (garrigue, oak woodland, and mixed oak/pine woodland) in Lebanon. Visitation rates to flowers were extremely low and restricted to male bees. The reproductive success of O. galilaea under ambient conditions was 29.3% (±2.4), compared to 89.0% (±2.1) in plants receiving cross-pollination by hand. No difference in reproductive success was found between habitat types, but values of reproductive success were positively correlated to the abundance of male bees. Pollination limitation can have negative impacts on the population growth of orchids, and this study provides clear evidence for more holistic approaches to habitat conservation to support specific interactions.