Low genetic variation reduces cross-compatibility and offspring fitness in populations of a narrow endemic plant with a self-incompatibility system

Genetic variation was shown earlier to bereduced in smaller populations of the narrowendemic putatively self-incompatible Cochlearia bavarica. To test whether thisnegatively affects plant fitness by reducedavailability of compatible mates and byinbreeding depression, we studied effects ofpopulation size and pollination treatments oncross-compatibility and offspring fitness in 16isolated populations of this plant. After openpollination, compatibility of crosses (i.e.,whether at least one fruit developed per markedflower), fruit set of compatible crosses, andcumulative fitness (number of plants permaternal ovule) after 14 months in a commongarden were lower for plants from smallerpopulations. Throughout the study, cumulativefitness was lower after hand pollination withpollen of one donor than after open pollination(finally 73.4% lower), suggesting that severalpollen donors or single pollen donors of higherquality are involved in open pollination.Moreover, cumulative fitness was lower afterhand selfing than after hand outcrossing(finally 69.4% lower), indicating bothinbreeding depression and reduced compatibilityafter selfing. High self-compatibility(40.6%), dry stigmas, and differences in thecompatibility of 11 of 33 experimentalreciprocal crosses between plant pairsconfirmed that C. bavarica has asporophytic self-incompatibility system, as iscommon in the Brassicaceae. Our studydemonstrates, that plants in smallerpopulations of species with a sporophyticself-incompatibility system can experiencetwofold fitness reductions associated withreduced genetic variability, i.e., twofoldgenetic Allee effects: via reducedcross-compatibility and via reduced offspringfitness.     

Relationship between population size, allozyme variation, and plant performance in the narrow endemic Cochlearia bavarica

We studied population viability in relation topopulation size and allelic variation in thenarrowly-endemic, monocarpic perennial plantCochlearia bavarica in Bavaria. In 1996,we analysed allelic variation by allozymeelectrophoresis in 24 populations ranging from8–2000 flowering individuals. Fitness-relatedcharacters were investigated in 22 of the 24populations in the field in 1996 (reproductiveand vegetative traits) and 1998 (reproductivetraits only). Differences in allozyme patternwere large between a south-eastern and awestern population group. Genetic diversity,assessed by the Shannon-Wiener diversity index,was low within but high among populations.Small populations had fewer alleles per locus,fewer polymorphic loci, lower observedheterozygosity, and lower genetic diversitythan large populations. Environmentalvariables were not significantly correlatedwith population size or fitness with theexception of light availability, indicatingthat habitat quality was similar for large andsmall populations. Population size showedpositive correlations with number of flowers,fruit set per plant, number of seeds per fruit,and total seed output per plant. Fruit set andnumber of seeds per fruit were positivelycorrelated with the observed heterozygosity andthe proportion of polymorphic loci. We usedpath analyses to study the possible causalrelationships among population size, allelicvariation, and reproductive characters. Thesemodels showed that allelic variation had nodirect influence on reproductive characters,whereas population size did. We conclude thatat present population size reduces viabilityand also reduces allelic variation; but thereduced allelic variation may in the longerterm have negative feed-backs on bothpopulation size and viability.     

Pollen limitation and variation in floral longevity in gynodioecious Potentilla tanacetifolia

Pollination limitation is common in flowering plants and is thought to be a factor driving the evolution of floral traits.The plasticity of floral longevity to pollination may be an adaptation of plants to pollen limitation.However,this adaptation is less critical in short-lived flowers.To evaluate pollen limitation and the plasticity of floral longevity to pollination in Potentilla tanacetifolia,a gynodioecious herb with short-lived flowers,we analyzed its breeding system,tested sex-differential pollen limitation,and compared variations in floral display size in natural populations in Duolun County,Inner Mongolia,China.Hand pollination experiments and pollinator exclusion treatments revealed that P tanacetifolia is self-compatible and non-autonomously apomictic and shows sex-differential pollen limitation.The plasticity of floral longevity to pollination was observed; the floral duration of female plants was prolonged by approximately 3-4 hours with pollination exclusion treatment.Moreover,the percentage of flowers displayed on female plants during pollination exclusion treatment was significantly higher than that during natural pollination.Under natural pollination conditions,the percentage of flowers displayed on female plants was significantly higher than on hermaphrodite plants.Furthermore,approximately 50％ of the pollen grains spread out of the anthers of hermaphrodite flowers within 2 h of anthesis; the number of pollen grains adhering to the stigmas of hermaphrodite flowers was significantly higher than that adhering to female flowers when flowers shed their petals.These results indicate that variation in floral longevity may be an adaptive strategy to pollination conditions for gynodioecious P tanacetifolia.     

Nitrogen limitation in dryland ecosystems: Responses to geographical and temporal variation in precipitation

We investigated the relationship between plant nitrogen limitation and water availability in dryland ecosystems. We tested the hypothesis that at lower levels of annual precipitation, aboveground net primary productivity (ANPP) is limited primarily by water whereas at higher levels of precipitation, it is limited primarily by nitrogen. Using a literature survey of fertilization experiments in arid, semi-arid, and subhumid ecosystems, we investigated the response of ANPP to nitrogen addition as a function of variation in precipitation across geographic gradients, as well as across year-to-year variation in precipitation within sites. We used four different indices to assess the degree of N limitation: (1) Absolute Increase of plant production in response to fertilization (the slope of ANPP vs. amount of added N at different levels of annual precipitation); (2) Relative Response (the percent increase in fertilized over control ANPP at different levels of N addition); (3) Fertilizer Use Efficiency (FUE, the absolute gain in productivity per amount of fertilizer N), and (4) Maximum Response (the greatest absolute increase in ANPP at saturating levels of N addition). Relative Response to fertilization did not significantly increase with increasing precipitation either across the geographic gradient or across year-to-year variation within sites. Nor did the Maximum Response to fertilization increase with increasing precipitation across the geographic gradient. On the other hand, there was a significant increase in the Absolute Increase and FUE indices with both geographical and temporal variation in precipitation. Together, these results indicate that there is not necessarily a shift of primary limitation from water to N across the geographic water availability gradient. Instead, our results support the hypothesis of co-limitation. The apparently contradictory results from the four indices of N limitation can best be explained by an integration of plant ecophysiological, community, and ecosystem mechanisms whereby plants are co-limited by multiple resources, species shifts occur in response to changing resource levels, and nitrogen and water availability are tightly linked through biogeochemical feedbacks.     

Pollen and water limitation in Astragalus scaphoides, a plant that flowers in alternate years

Mast seeding is common in plant populations, but its causes have rarely been tested experimentally. We tested mechanisms of alternate-year flowering and fruit set in an iteroparous, bee-pollinated, herbaceous plant, Astragalus scaphoides, in semi-arid sagebrush steppe. Patterns of reproduction from 1986 to 1999 indicated that spring precipitation was a cue for synchronous flowering, and that increased pollination in high-flowering years was a fitness advantage of synchrony. We tested these patterns by adding supplemental water and pollen to plants in high- and low-flowering sites and years. Supplemental water had no effect on flowering or seed set, so water is not a proximate cue for reproduction, though it could be important over longer (>3 year) time scales. Supplemental pollination increased fruit set in low- but not high-flowering years, indicating that synchronous flowering increases pollination success. Many shorter-term studies also report increased fruit set after pollen supplementation, but not after resource addition. This pattern may reflect the fact that plants can store and reallocate resources, but not pollen, across multiple years. For animal-pollinated herbs such as these, uniting theories about pollination ecology and mast seeding may promote an understanding of the mechanisms that determine patterns of reproduction over time.     

Pollen and resource limitation in a gynodioecious species

Differences between plant sex morphs in pollen or resource availability may affect their relative fitness and thereby the sex ratio of dimorphic species. In gynodioecious species, in which hermaphroditic and female plants coexist, a variety of factors (e.g., hermaphrodite self-fertility or rarity or pollinator discrimination against females) might be expected to lead to stronger pollen limitation in females than in hermaphrodites. On the other hand, females have been found to be superior compared to hermaphrodites in low-nutrient conditions. The effects of supplemental hand-pollination and resource addition on the reproductive output of the self-fertile gynodioecious perennial Geranium sylvaticum (Geraniaceae) were tested for several populations that differ in their female frequency (4.4-23.0%). Both pollen and resource availability limited fruit set and the number of seeds produced per plant; however, seed set (i.e., the number of seeds produced per fruit) was limited only by resources. Because pollen limitation in females did not correlate with female frequency, our results suggest that pollen limitation in females does not depend on the frequency of the pollen-producing hermaphrodites. Furthermore, because pollen and resource availability limited reproductive output of both sex morphs, these factors may not contribute significantly to maintenance and evolution of gynodioecy in G. sylvaticum.     

The biology and ecology of narrow endemic and widespread plants: a comparative study of trait variation in 20 congeneric pairs

The objective of this study is to examine whether habitat, herbivory and traits related to resource acquisition, resource conservation, reproduction and dispersal differ between narrow endemic plant species and their widespread congeners. We undertook pairwise contrasts of 25 ecological characteristics and biological traits in 20 congeneric pairs of narrow endemic and widespread plant species in the French Mediterranean region. Within each pair, the two species had the same life-form, pollination mode and dispersal mode. Endemic species differed significantly from widespread congeners for a number of attributes. Endemic species occur in habitats on steeper slopes, with higher rock cover and in lower and more open vegetation than their widespread congeners. Endemic species are significantly smaller than widespread species, but show no differences in traits related to resource acquisition (specific leaf area, leaf nitrogen content, maximum photosynthetic rate) or resource conservation (leaf dry matter content). After accounting for their smaller stature, endemic species produce fewer and smaller flowers with less stigma-anther separation and lower pollen/ovule ratios and produce fewer seeds per plant than their widespread congeners. No consistent variation in seed mass and propagule structure was found between congeneric species. Herbivory levels did not differ between congeneric species. Ecological characteristics, notably the occupation of rocky habitats with low aboveground competition, may thus have played an important role in the differentiation of narrow endemic species in the Western Mediterranean. Morphological and ecophysiological traits of narrow endemic species indicate that they are not more stress-tolerant than their widespread congeners. Lower investment in pollen transfer and seed production suggest that local persistence is a key feature of the population ecology of narrow endemic species.     

Pollen limitation meets resource allocation: towards a comprehensive methodology

The standard method of measuring pollen limitation is to add pollen to a number of flowers, preferably to a whole plant, and to compare fruit and seed set with that of naturally pollinated flowers on other plants. In 25 yr of research, this method has yielded valuable data, but it is difficult to use in large plants. This has caused a bias in the available data towards smaller, herbaceous plants with relatively few flowers. I argue that, in order to widen our knowledge of how pollen limitation affects plants, we should go beyond whole-plant pollen addition and change our concept of how a flowering plant functions. The traditional method does not take into account the variation in and dynamics of resource allocation and pollen availability. The concept of integrated physiological units (IPUs) does, but, although it has been applied to pollination biology, it has not received the attention it deserves. I use this article to present its merits again, to propose a step-by-step methodology for studying pollen limitation, and to examine factors influencing possible plant strategies.     

Genetic and ecological consequences of recent habitat fragmentation in a narrow endemic plant species within an urban context

Understanding the timescales that shape spatial genetic structure is pivotal to ascertain the impact of habitat fragmentation on the genetic diversity and reproductive viability of long-lived plant populations. Combining genetic and ecological information with current and past fragmentation conditions allows the identification of the main drivers important in shaping population structure and declines in reproduction, which is crucial for informing conservation strategies. Using historic aerial photographs, we defined the past fragmentation conditions for the shrub Conospermum undulatum, a species now completely embedded in an urban area. We explored the impact of current and past conditions on its genetic layout and assessed the effects of genetic and environmental factors on its reproduction. The historically high structural connectivity was evident in the genetics of the species. Despite the current intense fragmentation, we found similar levels of genetic diversity across populations and a weak spatial genetic structure. Historical connectivity was negatively associated with genetic differentiation among populations and positively related to within-population genetic diversity. Variation partitioning of reproductive performance explained?~?66% of the variance, showing significant influences for genetic (9%), environmental (15%), and combined (42%) fractions. Our study highlights the importance of considering the historical habitat dynamics when investigating fragmentation consequences in long-lived plants. A detailed characterization of fragmentation from 1953 has shown how low levels of genetic fixation are due to extensive gene flow through the non-fragmented landscape. Moreover, knowledge of the relationships between genetic and environmental variation and reproduction can help to implement effective conservation strategies, particularly in highly dynamic landscapes.

     

Environmental factors driving arbuscular mycorrhizal fungal communities associated with endemic woody plant Picconiaazorica on native forest of Azores

Arbuscular mycorrhizal fungi (AMF) play important key roles in the soil ecosystems as they link plants to the root-inaccessible part of soil. The aims of this study were to investigate which environmental factors influence the spatial and temporal structuring of AMF communities associated to Picconia azorica in two Azorean islands (Terceira and São Miguel islands), and investigate the seasonal variation in AMF communities between the two islands. Communities of AMF associated with P. azorica in native forest of two Azorean islands (Terceira and São Miguel) were characterised by spore morphology or molecular analysis. Forty-five AMF spore morphotypes were detected from the four fragments of P. azorica forest representing nine families of AMF. Acaulosporaceae (14) and Glomeraceae (9) were the most abundant families. AMF density and root colonisation varied significantly between islands and sampling sites. Root colonisation and spore density exhibited temporal patterns, which peaked in spring and were higher in Terceira than in São Miguel. The relative contribution of environmental factors showed that factors such as elevation, relative air humidity, soil pH, and soil available P, K, and Mg influenced AMF spore production and root colonisation. Different sporulation patterns exhibited by the members of the commonest families suggested different life strategies. Adaptation to a particular climatic and soil condition and host phenology may explain seasonal differences in sporulation patterns. Cohorts of AMF associated to P. azorica are shaped by regional processes including environmental filters such as soil properties and natural disturbance.     

Correction to: Genetic and ecological consequences of recent habitat fragmentation in a narrow endemic plant species within an urban context

Biodiversity and Conservation -     

Seed and microsite limitation in Rheum nobile,a rare and endemic plant from the subnival zone of Sino-Himalaya

Background: Species persistence, particularly in monocarpic species, depends on the successful recruitment of individuals. An understanding of the factors that limit the recruitment of rare monocarpic plant species is therefore vital for their conservation.

Aims: To identify the factors limiting the recruitment of Rheum nobile, a rare and highly specialised monocarpic giant herb endemic to the high eastern Himalayas.

Methods: Seed sowing (seeds added or not added) and seedling transplanting experiments were conducted in disturbed (vegetation removed) and undisturbed plots in the vicinity of established populations of R. nobile to explore the mechanisms of recruitment limitation. Four levels of photosynthetically active radiation (0, 15, 30 and 50 μmol m^?2 s^?1) and two sowing positions (beneath and above grass litter or moss layer) were manipulated in the laboratory to determine how ground cover limited seedling emergence.

Results: Seed addition increased seedling recruitment. Disturbance significantly increased seedling emergence and establishment. Seed germination significantly decreased with the reduction of light availability, but 31.7% of all seeds germinated in complete darkness. Seedling emergence was close to zero when seeds were sown on top of a layer of grass litter or moss, but rose to 34.5% when the seeds were sown beneath such layers.

Conclusions: Our results indicate that the recruitment of R. nobile is limited by a combination of seed and microsite availability. Therefore, in order to conserve this species, we suggest adding seeds to suitable sites and implementing soil disturbances in existing populations to create suitable microsites.     

ISSR variation in the endemic and endangered plant Cycas guizhouensis (Cycadaceae)

BACKGROUND AND AIMS: Cycas guizhouensis (Cycadaceae) is a rare and endangered species endemic to the southwest of China. An investigation was undertaken into the genetic variation of wild populations. METHODS: ISSR markers were used to determine the genetic variation within and between 12 extant populations of this species. KEY RESULTS: Low genetic diversity (at population level, P = 14.21 %, H(E) = 0.0597; at species level, P = 35.90 %, H(T) = 0.1082) and a high degree of differentiation among populations (G(ST) = 0.4321) were detected. CONCLUSIONS: This genetic structure is considered to be due to the combined effects of slow biochemical evolution, genetic drift, inbreeding and limited gene flow between populations. Based on these findings, strategies are proposed for the genetic conservation and management of the species.     

Pollen limitation of reproductive effort in willows

Summary Pollen limitation of seed set differs from resource limitation in its implications for the evolution of floral traits. Willow flowers attract insects, but also abundantly produce wind-dispersed pollen. I demonstrated pollen limitation in single branches bearing 2–4 inflorescences (catkins) in a field experiment with five species by artificially increasing or decreasing the pollen load. Because the responses by single branches might be explained by diversion of resources to better-pollinated branches within a plant, a second experiment with one species tested both pollen limitation of whole plants and the autonomy of catkins. Seed set of single willow catkins is unaffected by experimental alterations of seed set in other catkins on the same plant. Hand-pollination of single catkins and of whole plants increased seed set to the same degree, suggesting there is little or no competition for resources between catkins only 5–10 cm apart. Thus, seed set in willows appears to be pollen limited, favoring insect pollination and the evolution of entomophilous traits. The data support previous views that willows have a dual pollination system utilizing wind and insects.     

Genetic variation within and among populations of the narrow endemic,delphinium viridescens (Ranunculaceae)

Delphinium viridescens Leiberg (Ranunculaceae) is a narrow endemic, known from only 17 populations in the Wenatchee Mountains of central Washington. Electrophoretic data for ten loci and approximately 50 plants from each of the 17 populations indicate high levels of variability both within and among populations. High levels of heterozygosity, coupled with a general conformance of genotype frequencies to Hardy-Weinberg expectations, suggest that D. viridescens is an outcrossing species, and outcrossing rates range from 0.391 to 1.454. Although all populations share a suite of common alleles, interpopulational differences in allele frequency were detected. Nei's genetic identities range from 0.883 to 1.000, with a mean of 0.964, suggesting only slight differentiation among populations and at least moderate levels of interpopulational gene flow. However, F_ST values indicate greater interpopulational heterogeneity. The data from this study are intended for use in developing a management plan for D. viridescens, to maintain the viability of this species, and to assess the need for its listing as threatened or endangered.     

Number of endemic and native plant species in the Galápagos Archipelago in relation to geographical parameters

By simple and multiple regression analyses we investigate updated species numbers of endemic and native vascular plants and seed plants in the Galápagos Archipelago in relation to geographical parameters. We find that the best models to describe species numbers are regression models with log-transformed species numbers as dependent and log-transformed modified area (i.e. area not covered with barren lava) as an independent variable. This holds both for total species number, for native species number, for endemic species number and for total number of seed plants as well as number of endemic seed plants. For the ratio between endemic and native species, modified area is also the major significant variable, but with a negative regression slope.
Multiple regression models show that some isolation measures are significant contributors and may explain some of the residual variation, but their contribution to total explained variation is in general small.
The results show that the species area relationships are different for native and endemic species. This is discussed in relation to classical island biogeographical models, and the concepts of radiative speciation.     

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.     

Allozyme variation of the endangered endemic plant Myricaria laxiflora: Implications for conservation

Myricaria laxiflora, a riparian plant that naturally occurs in the riverbanks of the Yangtze River Valley, has become extinct across its entire geographical distribution range in the wild due to the construction of the Three Gorges Dam. The allozyme variation of M. laxiflora populations was investigated in the present study. Mean number of alleles per locus (A) was 2.35, and the observed heterozygosity (H_o) and expected heterozygosity (H_e) were 0.35 and 0.30, respectively. Six populations showed significant excesses of heterozygotes based on the examination of a multilocus fixation index (F_IS). The population genetic divergence of M. laxiflora is high (G_ST = 0.144 and ?_B = 0.131) and the analysis of molecular variance analysis shows that 19.71% of the total genetic variation is caused by the difference between populations. Based on the obtained genetic information, six management units have been identified, all of which are expected to enhance the effective management of the remaining and transplanted individuals of this endangered species in the future.     

Pollen limitation in a single year is not compensated by future reproduction

Oecologia - Seed production is critical to the persistence of most flowering plant populations, but may be strongly pollen limited. To what extent long-lived plants can compensate pollen limitation...     

Patterns of abundance of a narrow endemic species in a tropical and infertile montane habitat

We examined the entire spatial distribution of a narrow endemic shrub (Coccoloba cereifera, Polygonaceae) in Serra do Cipó, Brazil. We tested the hypothesis that a narrow endemic species would show a gradual decline in either size and density towards the edges of its distribution. The contribution of soil specificity and post-fire growth to C. cereifera abundance and distribution were also investigated. C. cereifera showed multimodal and highly aggregated distribution pattern at several scales, from 25 m² to 3000 m² (blocked quadrat variance analyses). This pattern seems to be strongly related to the predominance of clonal recruitment and to the close association of the species to sandfields, which have discrete distribution between gallery forests and rocky outcrops. Population density did not decline towards the edge of the species distribution. Plants near the distribution boundaries had slightly more leaves and more inflorescences per plant (p<0.005), but there was no significant change in the mean number of ramets per clone. The absence of large plants in some populations at the center of the species distribution may be related to the higher frequency of fire in this region, killing aerial plant parts. Nearly all aggregations had inverse-J shaped size-distribution, suggesting effective recruitment of ramets, most frequently via asexual reproduction. Similar patterns of plant abundance may be common in fire-prone habitats characterized by infertile, and well-drained soils since these areas generally have high numbers of endemic plants, with strong soil specificity. Possible mechanisms for the observed pattern are discussed considering current models concerning distribution of abundance of species.