Recurrent fruit harvesting reduces seedling density but increases the frequency of clonal reproduction in a tropical tree

Studies on the ecological impacts of non‐timber forest products (NTFP) harvest reveal that plants are often more resilient to fruit and seed harvest than to bark and root harvest. Several studies indicate that sustainable fruit harvesting limits can be set very high (>80% fruit harvesting intensity). For species with clonal and sexual reproduction, understanding how fruit harvest affects clonal reproduction can shed light on the genetic risks and sustainability of NTFP harvest. We studied 18 populations of a gallery forest tree, Pentadesma butyracea (Clusiaceae), to test the impact of fruits harvest, climate and habitat size (gallery forest width) on the frequency of sexual or clonal recruitment in Benin, West Africa. We sampled populations in two ecological regions (Sudanian and Sudano‐Guinean) and in each region, we selected sites with low, moderate and high fruit harvesting intensities. These populations were selected in gallery forests with varying width to sample the natural variation in P. butyracea habitat size. Heavily harvested populations produced significantly less seedlings but had the highest density and proportion of clonal offspring. Our study suggests that for plant species with dual reproductive strategy (via seeds and clonal), fruit harvesting and associated disturbances that come with it can lead to an increase in the proportion of clonal offspring. This raises the issue that excessive fruit harvest by increasing the proportion of clonal offspring to the detriment of seed originated offspring may lead to a reduction in genetic diversity with consequence on harvested species capability to withstand environmental stochasticity.     

Accessory costs of seed production

Accessory costs of reproduction are those that are necessary to mature a seed, but that do not involve the direct cost of provisioning the seed itself. This study aims to quantify accessory costs in a range of species, and test whether they decrease as a proportion of total reproductive expenditure with increasing seed mass, as might be expected if economies of scale came into play at larger seed sizes. We also test whether accessory costs varied with growth form, pollination mode, and dispersal mode, with the expectation that biotic pollination and dispersal modes should incur greater costs. Reproductive allocation (dry biomass) over one season, was calculated for 14 diclinous angiosperm species. Accessory costs averaged 73% of total reproductive allocation, with the majority spent on packaging and dispersal. Total accessory costs, packaging and dispersal costs, and costs incurred prior to pollination were proportional to direct costs of reproduction in major axis regressions. However, larger seeded species incurred significantly greater costs associated with aborted seeds and fruits, and matured a smaller proportion of ovules. This is consistent with larger seeded species being more selective of the ovules/embryos matured than small-seeded species. Total accessory costs, and proportion of ovules aborted, were also significantly greater for biotically dispersed species, but only due to an association with larger seed masses. Costs associated with abortions were lower for biotically pollinated species, due to a general trend of more ovules per ovary, resulting in greater cost sharing. This study demonstrates that expenditure on items other than seeds accounts for the majority of reproductive allocation in flowering plants. Yet, far more literature exists on seed mass variation than on investment in accessory structures. We found a proportional relationship between accessory costs and seed mass that warrants further investigation within the context of selection on margin returns on investment.     

Population dynamics of the climate‐sensitive endangered perennial Ferula sadleriana Ledeb. (Apiaceae)

Ancient plant species surviving in isolated small populations are particularly vulnerable to extinction, therefore understanding their population dynamics is necessary for conservation. The iteroparous perennial relic endemic Ferula sadleriana Ledeb. (Apiaceae) is restricted to seven distant localities in the Carpathian Basin, where it inhabits rocky hills. We monitored the species' largest population on the Pilis Hill, Hungary, over 14–19 years (depending on trait) between 1979 and 2010, and relationships were sought between climatic properties and population attributes. The population of 4000 ± 1509 emergent individuals underwent large interannual fluctuations, with the vegetative stage displaying sevenfold and the reproductive stage twenty‐eight‐fold differences. Spring and early summer precipitation had a marked influence on abundances and seed set. Alternating years of high and low counts of reproductive plants suggest costs of reproduction that most probably incur prolonged dormancy and retrogression to the vegetative stage. Seed set was positively influenced by number of reproductive plants over years and by plant size within a year. Ungulates nullify yearly reproductive output by grazing on reproductive individuals. This is particularly intense in dry summers, when reproductive output is already low. The strong precipitation response of abundance, absence of clonal propagation and soil seed bank, and geographical isolation of the populations place F. sadleriana at considerable risk under an increasingly variable and extreme climate. Management should seek to maintain the species' original habitat mosaic (potentially compensating for climate variation), minimize grazing damage and anthropogenic disturbance, and establish ex situ conservation programs to provide propagules for eventual reintroduction.     

Effect of habitat area and isolation on plant trait distribution in European forests and grasslands

A number of studies show contrasting results in how plant species with specific life‐history strategies respond to fragmentation, but a general analysis on whether traits affect plant species occurrences in relation to habitat area and isolation has not been performed. We used published data from forests and grasslands in north‐central Europe to analyse if there are general patterns of sensitivity to isolation and dependency of area for species using three traits: life‐span, clonality, and seed weight. We show that a larger share of all forest species was affected by habitat isolation and area as compared to grassland species. Persistence‐related traits, life‐span and clonality, were associated to habitat area and the dispersal and recruitment related trait, seed weight, to isolation in both forest and grassland patches. Occurrence of clonal plant species decreased with habitat area, opposite to non‐clonal plant species, and long‐lived plant species decreased with grassland area. The directions of these responses partly challenge some earlier views, suggesting that further decrease in habitat area will lead to a change in plant species community composition, towards relatively fewer clonal and long‐lived plants with large seeds in small forest patches and fewer clonal plants with small seeds in small grassland patches. It is likely that this altered community has been reached in many fragmented European landscapes consisting of small and isolated natural and semi‐natural patches, where many non‐clonal and short‐lived species have already disappeared. Our study based on a large‐scale dataset reveals general and useful insights concerning area and isolation effects on plant species composition that can improve the outcome of conservation and restoration efforts of plant communities in rural landscapes.     

Potential ‘costs of reproduction’ in a skink: Inter‐ and intrapopulational variation

Reproductive costs are important determinants of reproductive effort in squamate reptiles. Consequently, differences in costs of reproduction between populations of geographically or climatically widespread species are likely to result in different patterns of reproductive effort. In the present study, the effect of pregnancy on sprint speed was examined in a small viviparous skink, Niveoscincus ocellatus (Gray 1845), from two populations at the climatic extremes of its distribution. Decreased sprint speed has the potential to be an important cost of reproduction in this species, through a reduced ability to avoid predation and/or decreased foraging efficiency. Lizards inhabiting the colder site were larger than those from the warmer site and, contrary to predictions from life history theory, had a higher reproductive effort. In both populations, sprint speed was lower in pregnant lizards than in either the same individuals after birth or non‐pregnant control lizards. Within each population, sprint speed was unrelated to the level of reproductive effort of the female in terms of either absolute mass of the reproductive burden or the burden relative to her post‐partum body mass. However, within each population, the mass of the clutch that an individual female was carrying relative to snout–vent length was an important determinant of her sprint speed while pregnant. Thus, within each population, a relatively high reproductive burden may potentially increase costs of reproduction in this species. Despite this relationship and predictions from life history theory suggesting that annual reproductive effort will be lower in populations with a large body size and delayed maturity, it is suggested that a higher reproductive effort at the cold site is possible because they have a higher absolute sprint speed because of their larger size and a relatively higher abundance of cover at the cold site, and differences in predation pressure may alter selective pressures on reproductive investment.     

Differential costs of reproduction in females and hermaphrodites in a gynodioecious plant

Background and Aims

Plants exhibit a variety of reproductive systems where unisexual (females or males) morphs coexist with hermaphrodites. The maintenance of dimorphic and polymorphic reproductive systems may be problematic. For example, to coexist with hermaphrodites the females of gynodioecious species have to compensate for the lack of male function. In our study species, Geranium sylvaticum, a perennial gynodioecious herb, the relative seed fitness advantage of females varies significantly between years within populations as well as among populations. Differences in reproductive investment between females and hermaphrodites may lead to differences in future survival, growth and reproductive success, i.e. to differential costs of reproduction. Since females of this species produce more seeds, higher costs of reproduction in females than in hermaphrodites were expected. Due to the higher costs of reproduction, the yearly variation in reproductive output of females might be more pronounced than that of hermaphrodites.

Methods

Using supplemental hand-pollination of females and hermaphrodites of G. sylvaticum we examined if increased reproductive output leads to differential costs of reproduction in terms of survival, probability of flowering, and seed production in the following year.

Key Results

Experimentally increased reproductive output had differential effects on the reproduction of females and hermaphrodites. In hermaphrodites, the probability of flowering decreased significantly in the following year, whereas in females the costs were expressed in terms of decreased future seed production.

Conclusions

When combining the probability of flowering and seed production per plant to estimate the multiplicative change in fitness, female plants showed a 56 % and hermaphrodites showed a 39 % decrease in fitness due to experimentally increased reproduction. Therefore, in total, female plants seem to be more sensitive to the cost of reproduction in terms of seed fitness than hermaphrodites.     

Phenotypic plasticity and specialization in clonal versus non-clonal plants: A data synthesis

Reproductive strategies can be associated with ecological specialization and generalization. Clonal plants produce lineages adapted to the maternal habitat that can lead to specialization. However, clonal plants frequently display high phenotypic plasticity (e.g. clonal foraging for resources), factors linked to ecological generalization. Alternately, sexual reproduction can be associated with generalization via increasing genetic variation or specialization through rapid adaptive evolution. Moreover, specializing to high or low quality habitats can determine how phenotypic plasticity is expressed in plants. The specialization hypothesis predicts that specialization to good environments results in high performance trait plasticity and specialization to bad environments results in low performance trait plasticity. The interplay between reproductive strategies, phenotypic plasticity, and ecological specialization is important for understanding how plants adapt to variable environments. However, we currently have a poor understanding of these relationships. In this study, we addressed following questions: 1) Is there a relationship between phenotypic plasticity, specialization, and reproductive strategies in plants? 2) Do good habitat specialists express greater performance trait plasticity than bad habitat specialists? We searched the literature for studies examining plasticity for performance traits and functional traits in clonal and non-clonal plant species from different habitat types. We found that non-clonal (obligate sexual) plants expressed greater performance trait plasticity and functional trait plasticity than clonal plants. That is, non-clonal plants exhibited a specialist strategy where they perform well only in a limited range of habitats. Clonal plants expressed less performance loss across habitats and a more generalist strategy. In addition, specialization to good habitats did not result in greater performance trait plasticity. This result was contrary to the predictions of the specialization hypothesis. Overall, reproductive strategies are associated with ecological specialization or generalization through phenotypic plasticity. While specialization is common in plant populations, the evolution of specialization does not control the nature of phenotypic plasticity as predicted under the specialization hypothesis.     

Floral display and habitat quality affect cost of reproduction in Primula farinosa

Costs of reproduction should depend on resource availability and on reproductive effort, which in turn may depend on traits influencing reproductive success. Therefore, variation in both habitat quality and reproductive traits should be considered when assessing reproductive costs. We investigated the effect of habitat quality and floral display on the costs of reproduction in the perennial herb Primula farinosa. In the study area, P. farinosa occurs in habitats that differ in water availability, which strongly influences plant performance. Furthermore, it displays a scape length dimorphism, with two distinct scape morphs differing in attractiveness to pollinators and reproductive success. To test the hypothesis that the cost of fruit production is higher in the long‐scaped than in the short‐scaped morph, and depends on water availability, we manipulated reproductive investment in eight P. farinosa populations along a gradient of soil moisture. Fruit set was higher in long‐scaped individuals, and prevention of fruit set increased flower production in the following year among long‐scaped, but not among short‐scaped plants. Furthermore, costs of fruit production were evident at low and high moisture levels but not at intermediate levels. The results demonstrate an association between a genetically determined difference in floral display and cost of reproduction, and suggest that costs of reproduction are non‐linearly related to water availability. They thus indicate links between the evolution of plant reproductive traits and plant life histories, and between habitat quality and optimal life history.     

Effects of diapause on post‐diapause reproductive investment in the moth Ostrinia scapulalis

Diapause is a strategy used by many insect species to survive adverse environmental conditions. However, diapause incurs costs that may have adverse effects on post‐diapause development and reproduction. We herein investigated the effects of diapause on the post‐diapause reproductive investment of males and females in a multivoltine moth, the adzuki bean borer, Ostrinia scapulalis (Walker) (Lepidoptera: Crambidae). We found that (1) post‐diapause males and females were smaller and had lower mating success than non‐diapause individuals, (2) post‐diapause females had lower fecundity and shorter longevity than non‐diapause females, (3) post‐diapause males transferred similar numbers of eupyrene and apyrene sperm as non‐diapause males, (4) the fecundity and longevity of non‐diapause females mated with post‐diapause males and those mated with non‐diapause males were not significantly different, and (5) no significant relationship was found between diapause duration (short and long) and post‐diapause reproductive investments in both males and females. These results suggest that post‐diapause males did not reduce reproductive investment in spite of the cost of diapause, and the significant decline in reproductive output in post‐diapause females was due to their reduced body weight and longevity, which appeared to be direct consequences of the cost of diapause.     

Indirect positive effects of a parasitic plant on host pollination and seed dispersal

Parasitic plants often have a strong fitness‐impact on their plant hosts through increased host mortality and reduced or complete suppression of reproduction. Tristerix corymbosus (Loranthaceae) is a hemiparasitic mistletoe that infects a wide range of host species along its distribution range. Among such species, Rhaphithamnus spinosus (Verbenaceae) is a frequent host with a flowering and fruiting season partially synchronized with mistletoe reproductive phenology. As parasitized hosts have, in principle, a larger flower display and fruit crop size than non‐parasitized hosts, we examined whether host and parasite reproductive synchrony make infected hosts more attractive for pollinators and seed dispersers than uninfected hosts. Our results showed that pollinator visit rates did not differ between parasitized and non‐parasitized hosts. Conversely, seed rain was higher in parasitized than non‐parasitized individuals. The number of seeds fallen under non‐parasitized plants was spatially associated with crop size, while parasitized plants did not show such association. Finally, the number of seedlings of R. spinosus was significantly larger near parasitized than non‐parasitized hosts. Our results suggest that the presence of the mistletoe might be responsible of the higher reproductive success showed by the parasitized fraction of R. spinosus. This effect, however, seems to be related to seed dispersal processes rather than pollination effects.     

Demographic consequences of greater clonal than sexual reproduction in Dicentra canadensis

Clonality is a widespread life history trait in flowering plants that may be essential for population persistence, especially in environments where sexual reproduction is unpredictable. Frequent clonal reproduction, however, could hinder sexual reproduction by spatially aggregating ramets that compete with seedlings and reduce inter‐genet pollination. Nevertheless, the role of clonality in relation to variable sexual reproduction in population dynamics is often overlooked. We combined population matrix models and pollination experiments to compare the demographic contributions of clonal and sexual reproduction in three Dicentra canadensis populations, one in a well‐forested landscape and two in isolated forest remnants. We constructed stage‐based transition matrices from 3 years of census data to evaluate annual population growth rates, λ. We used loop analysis to evaluate the relative contribution of different reproductive pathways to λ. Despite strong temporal and spatial variation in seed set, populations generally showed stable growth rates. Although we detected some pollen limitation of seed set, manipulative pollination treatments did not affect population growth rates. Clonal reproduction contributed significantly more than sexual reproduction to population growth in the forest remnants. Only at the well‐forested site did sexual reproduction contribute as much as clonal reproduction to population growth. Flowering plants were more likely to transition to a smaller size class with reduced reproductive potential in the following year than similarly sized nonflowering plants, suggesting energy trade‐offs between sexual and clonal reproduction at the individual level. Seed production had negligible effects on growth and tuber production of individual plants. Our results demonstrate that clonal reproduction is vital for population persistence in a system where sexual reproduction is unpredictable. The bias toward clonality may be driven by low fitness returns for resource investment in sexual reproduction at the individual level. However, chronic failure in sexual reproduction may exacerbate the imbalance between sexual and clonal reproduction and eventually lead to irreversible loss of sex in the population.     

The interplay of pollination,costs of reproduction and plant size in maternal fertility limitation in perennial <Emphasis Type="Italic">Paeonia officinalis</Emphasis>

Although several factors can limit female fertility in perennial plants, rarely have they been jointly studied in a single species over several years. In this study we experimentally manipulate seed production and simultaneously analyse the potential contribution of pollen limitation, costs of reproduction and plant size to variation in seed output over a 3-year period in the perennial herb Paeonia officinalis, in southern France. Since this rare species is threatened by forest closure in many sites we also examine the causes of female fertility variation in relation to habitat closure (open habitat vs. woodland). P. officinalis has a partial self-incompatibility system and only very low ability for autonomous self pollination in the absence of pollinators. However, supplementary pollination of individual plants in three consecutive years did not significantly increase seed production above natural levels. Forest closure was associated with a decline in ovule and seed production, which again was not due to pollen limitation since supplementary pollination had no significant effect on seed set in the woodland habitat. Comparison of the maternal fertility of plants which were previously excluded from reproduction with those which were hand pollinated to maximise seed set in two previous years produced no evidence that seed production in year three is limited by costs associated with prior reproduction. Likewise, flowering probability was not related to prior seed production but was however positively related to plant size. The absence of any influence of pollen limitation or prior reproduction on seed production suggests that sub-maximal seed production in long-lived perennial herbs may be part of a size-dependent strategy that maximises life-time seed production and fitness without compromising survival.     

Phenology and fecundity in 11 sympatric pioneer species of Macaranga (Euphorbiaceae)in Borneo

Reproductive traits of tropical tree species vary predictably in relation to successional stage, but this variation may be due to the species' phylogenetic histories rather than selective pressures imposed by regeneration requirements. Reproductive phenology, tree size at the onset of reproduction, and fecundity of 11 sympatric, closely related Macaranga species were studied to investigate within-species variation in reproductive traits in relation to resource availability, and among-species variation in relation to other life-history traits (shade tolerance, seed size and maximum tree size, H(max)) and consequently the requirements for forest-gap colonization. Nine species reproduced in synchronous episodes, and two species reproduced continuously over 32 mo. Episodic reproduction was most intense in 1992 following a severe drought. For several species, reproductive trees had greater light availability, lower fecundity in lower light levels, and lower growth rates than nonreproductive trees, reflecting resource-limited reproduction. Among species, H(max) was negatively correlated with shade tolerance and seed size. Tree size at the onset of reproduction and fecundity was strongly linked to this axis of life-history variation, but phenological pattern was not. Absolute tree size at the onset of reproduction was positively correlated with H(max) and negatively correlated with shade tolerance. Relative size at reproductive onset was not correlated with shade tolerance or H(max). Fecundity ranged four orders of magnitude among species and was correlated positively with H(max) and negatively with seed size and shade tolerance. The interrelationships among these reproductive and other life-history traits are strongly correlated with the species' requirements for gap colonization.     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.     

Breeding system, flower visitors and seedling survival of two endangered species of Helianthemum (Cistaceae)

BACKGROUND AND AIMS: Helianthemum marifolium and H. caput-felis are two endangered plant species of the western Mediterranean. Several aspects of the reproduction of both species were examined to determine whether their rarity could be related to factors causing reproductive limitation. METHODS: The flowering and fruiting phenology of both species in two non-sympatric island populations (Mallorca, Balearic Islands, western Mediterranean) were compared. Hand-pollination experiments were conducted to determine their fruit and seed production under different pollen sources. The composition of the pollinator assemblage and the effect of temporal variation and sun exposure on reproductive output and seedling survival were also investigated. KEY RESULTS: Flowering periods were longer for H. marifolium than for H. caput-felis in the populations studied. Helianthemum marifolium is mostly an outbreeder, i.e. fruit and seed set was three-fold higher when pollen came from other plants. In H. caput-felis, neither fruit nor seed set was affected by pollination treatments. Flower visitors were more diverse for H. caput-felis than for H. marifolium. In both species, most floral visits were made by hymenopterans. The total number of pollinator visits varied significantly between years, decreasing more than two-fold from 2001 to 2002, in both species. In agreement with its outbreeder character, variation in reproductive output between years was also observed in H. marifolium. It showed a 50 % decrease in fruit set in 2002, a pollinator-poor year. Finally, seedling survival increased three- to six-fold from 2001 to 2002. A correlation between seedling size and survival had also been detected. CONCLUSIONS: Reproductive limitations were detected for neither species (i.e. fruit and seed set, pollination service and seedling survival on natural populations). Hence, the increasing rarity of these two species is probably a direct result of the destruction of their habitat.     

Interspecific competition changes reproductive output but does not increase reproductive costs in a grassland perennial

In plants, it is hypothesized that allocation trade-offs may appear only when expenditures like seed production are high or external resources are scarce. In this study, we tested whether reproductive costs are more pronounced under enhanced interspecific competition.In a common garden, we investigated phenotypic correlations between sexual reproduction, clonal growth and storage structures in the grassland perennial, Succisa pratensis. During the past 50 years, habitats of this species have faced an expansion of clonal grasses that increase competition intensity. We simulated this process by growing five populations of Succisa from high- and low-production habitats with its clipped and non-clipped competitor, Agrostis capillaris. In addition, we experimentally removed flower heads of Succisa plants from one population grown with and without A. capillaris.We demonstrated costs of sexual reproduction by flower-head removal (resulting in increased plant size and relative allocation to belowground structures) but not by phenotypic correlations. We found no evidence that reproductive costs increase in a competitive environment and the opposite pattern was shown in both approaches used. However, high competition intensity reduced relative investment to flower-head production. In plants from low-production habitats, competition also reduced the absolute number of flower heads and belowground biomass as a result of smaller plant size. We assume that populations from low-production habitats are more prone to extinction as they have a reduced likelihood of local persistence and of escape to more suitable habitats during advancing succession.     

Reproductive success in fragmented habitats: do compatibility systems and pollination specialization matter?

Abstract. This paper explores whether plant breeding system and pollination specialization influence the reproductive response of plants to habitat fragmentation. It is meaningful for conservation to predict a plant species’ extinction risk. We found 25 studies in the literature assessing the effects of habitat fragmentation on either pollination or reproductive success of 46 plant species to answer the following questions: 1. Are pollination and reproductive success of self‐incompatible species more likely to decline with habitat fragmentation than the pollination and reproductive success of self‐compatible species? Although most of the species showed statistically significant negative effects, the pollination and reproduction of self‐incompatible species were as likely to decline with fragmentation as those of self‐compatible species. 2. Are pollination and reproductive success of specialist plants more affected than the pollination and reproduction of generalist plants? Comparisons of fragmentation‐related changes in pollination and reproductive success between specialists and generalists do not support the hypothesis that specialization in pollination increases the risk of plant extinction. 3. Can self‐incompatible species offset their expected higher vulnerability to fragmentation by being, on average, more pollination generalist than self‐compatible species? In a larger data set on 260 species, we did not find significant differences in either the mean number or frequency distribution of numbers of flower‐visiting species or orders between self‐compatible and self‐incompatible species. Our review suggests that no generalizations can be made on susceptibility to fragmentation based on compatibility system and pollination specialization.     

The evolutionary ecology of mast seeding

The past seven years have seen a revolution in understanding the causes of mast seeding In perennial plants. Before 1987, the two main theories were resource matching (i.e. plants vary their reproductive output to match variable resources) and predator satiation (i.e. losses to predators are reduced by varying the seed crop). Today, resource matching is restricted to a proximate role, and predator satiation is only one of many theories for the ultimate advantage of masting. Wind pollination, prediction of favourable years for seedling establishment, animal pollination, animal dispersal of fruits, high accessory costs of reproduction and large seed size have all been advanced as possible causes of masting. Of these, wind pollination, predator satiation and environmental prediction are important in a number of species, but the other theories have less support. In future, Important advances seem likely from quantifying synchrony within a population, and examining species with very constant reproduction between years.     

Populations do not become less genetically diverse or more differentiated towards the northern limit of the geographical range in clonal Vaccinium stamineum (Ericaceae)

Geographically peripheral populations are expected to exhibit lower genetic diversity and higher differentiation than central populations because of their smaller size and greater spatial isolation. In plants, a shift from sexual to clonal asexual reproduction may further reduce diversity and increase differentiation. Here, these predictions were tested by assaying 36 inter-simple sequence repeat (ISSR) polymorphisms in 21 populations of the woody, clonal plant Vaccinium stamineum in eastern North America, from the range center to its northern limit where it has 'threatened' status. Populations decline in frequency, but not size or sexual reproductive output, across the range. Within-population diversity did not decline towards range margins. Modest genetic differentiation among populations increased slightly towards range margins and in small populations with high clonal propagation and low seed production, although none of these trends was significant. Low seed production and high clonal propagation were not associated with large-scale clonal spread. By combining demographic and genetic data, this study determined that increased population isolation, rather than reduced population size, can account for the weak increase in genetic differentiation at range margins.     

Mutualisms matter: pollination rate limits the distribution of oil‐secreting orchids

There are at least two immediate reasons why it is important to determine the role of biotic interactions, such as pollination, in limiting species distribution ranges. Firstly, if range limits are imposed by biotic factors, current and future distribution ranges might not be constrained by climate. Secondly, if biotic interactions limit the distribution ranges of species, anthropogenic impacts on these interactions are likely to have a major effect on biodiversity. Here we test the role of pollination in limiting plant distributions by studying plant community assembly in a guild of 15 oil‐secreting orchids (Coryciinae) along a pollination gradient. In all members of the guild, seed production depends on pollination by the oil‐collecting bee Rediviva peringueyi (Melittidae). While the mode of aboveground reproduction is uniform across the guild, the orchid species differ widely in their capacity for belowground clonal reproduction through the formation of bulbils, and hence span a range of predicted dependence on pollination (and subsequent seed set) for population persistence. Pollination rate by R. peringueyi varied across the landscape from 0 to 98% of flowers pollinated. With decreasing pollination, species richness of the orchid guild declined, and species were lost by the successive deletion of the least clonal species. Thus, pollination is shown to act as a biotic filter, excluding non‐clonal species from pollinator‐poor communities. The findings are consistent with the idea that pollination mutualisms matter ecologically by limiting the distribution of non‐clonal plants. Conversely, the results suggest that clonality allows some plant species to escape from the range of their pollinators.