Ecological and evolutionary mechanisms for low seed: ovule ratios: need for a pluralistic approach?

Central to the ecology and evolution of a broad range of plants is understanding why they routinely have submaximal reproduction manifested as low seed : ovule and fruit : flower ratios. We know much less about the processes responsible for low seed : ovule ratios than we do for fruit : flower ratios. Current hypotheses for low seed : ovule ratios are largely drawn from those for fruit : flower ratios, including proximate (ecological) causes of pollen limitation, resource limitation, and pollen quality, as well as the ultimate (evolutionary) hypothesis of "bet hedging" on stochastic pollination. Yet, such mechanisms operating on fruit : flower ratios at the whole-plant level may not best explain low seed : ovule ratios at the individual-flower level. We tested each of these proximate and ultimate causes for low seed : ovule ratios using the specialized pollination mutualism between senita cacti (Pachycereus schottii) and senita moths (Upiga virescens). Seed : ovule ratios were consistently low (approximately 0.61). Such excess ovule production by senita likely has a strong genetic component given the significant differences among plants in ovule number and the consistency in ovule production by plants within and among flowering seasons. Excess ovule production and low seed : ovule ratios could not be explained by pollen limitation, resource limitation, pollen quality, or bet hedging. Nevertheless, phenotypic selection analyses did show significant selection gradients for increased ovule number, suggesting that other evolutionary processes may be responsible for excess ovule production and low seed : ovule ratios. In contrast, low fruit : flower ratios at the whole-plant level were explained by an apparent equilibrium between pollen and resource limitation. Thus, mechanisms responsible for low fruit : flower ratios at the whole-plant level are not necessarily in accord with those of low seed : ovule ratios at the individual-flower level. This suggests that we may need to adopt a more pluralistic approach to seed : ovule ratios and consider alternative hypotheses, including a greater array of proximate and ultimate causes. Initial results of this study suggest that floral allometry, selection on correlated floral traits, stigma clogging with pollen grains, and style clogging with pollen tubes may provide promising avenues for understanding low seed : ovule ratios.     

Experimental Demonstration of an Allee Effect in American Ginseng

Abstract: Harvesting of wild American ginseng (   Panax quinquefolius ) for the herbal trade has lowered natural population sizes. We tested for reproductive limitation due to small population size (a form of the Allee effect) by experimentally planting "natural" populations numbering 4, 16, and 64 using 4-year-old cultivated plants. Plant size traits and reproductive traits ( bud, flower, green fruit, and mature fruit) were recorded through the ensuing summer. Fruit production per flower and per plant increased in proportion to flowering population size (  p = 0.0063 and p = 0.0017, respectively), strongly suggesting that an Allee effect occurs in very small populations. The increase in fruit production was not explained by either plant or inflorescence size differences. Although population size-dependent pollination, through insufficient pollinator visitation rate or pollen transfer rate, seems the most likely cause of the observed effects, our limited observations of pollinators were not sufficient to demonstrate a change in pollination rates as a function of population size. Knowledge of the presence as well as the mechanism underlying this Allee effect may be especially useful for management and determination of minimum viable population size of the species in the wild.     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Population Size and Reproduction in Phlox pilosa

Abstract: We analyzed the relationships between population size and reproductive characteristics in the perennial prairie forb Phlox pilosa , an obligate outcrossing butterfly-pollinated species. We examined 27 populations ranging in size from 9 to over 75,000 flowering ramets in two regions of the state of Iowa (eastcentral and northwest) in 1993 and 1994. We collected flowers from each population and scored them for pollen arrival to stigmas and number of pollen tubes. We collected fruiting ramets from each population at the end of the Phlox growing season and scored them for height, biomass, and reproductive variables, including the number of flowers initiated and opened and the number of capsules initiated and matured. In both years, population size was significantly correlated with the number of capsules matured per ramet. Differences between populations in capsule production were set primarily at the pollination stage. In 1993, pollen arrival to stigmas was significantly lower than in 1994 and was correlated with population size in eastcentral Iowa populations, indicating that lower reproduction in small populations that year was at least partially due to inadequate amounts of pollen being moved. In 1993, weather conditions likely depressed pollinator activity, but absolute capsule formation was high because of high flower production per ramet and high population densities. In 1994, when pollen arrival to stigmas was relatively high and unrelated to population size, outcross pollen movement was greater in larger populations. Increased efficacy of outcross pollen movement in 1994 may have resulted from lower flower production and less dense populations forcing greater pollinator movement between ramets or from variation between years in fine-scale spatial genetic substructuring of populations. Our results indicate that the viability of Phlox pilosa can be best ensured by protecting and creating populations of at least 1000–2000 flowering ramets.     

Scale dependence of reproductive failure in fragmented Echinacea populations

I investigated reproduction in a three-year study of Echinacea angustifolia, purple coneflower, growing in a fragmented prairie landscape. I quantified the local abundance of flowering conspecifics at individual-based spatial scales and at a population-based spatial scale. Regression analyses revealed that pollen limitation increased while seed set and fecundity decreased with isolation of individual plants. Isolation, defined as the distance to the k(th) nearest flowering conspecific, was a good predictor of pollen limitation, for all nearest neighbors considered (k = 1-33), but the strength of the relationship, as quantified by R2, peaked at intermediate scales (k = 2-18). The relationship of isolation to seed set and fecundity was similarly strongest at intermediate scales (k = 3-4). The scale dependence of individual density effects on reproduction (density of flowering plants within x meters) resembled that of isolation. Analyses at a population-based scale showed that pollen limitation declined significantly with population size. Seed set and fecundity also declined with population size, but significantly so only in 1998. Whether quantifying local abundance with population- or individual-based measures, reproductive failure due to pollen limitation is a consistent consequence of Echinacea scarcity. However, individual-based measures of local abundance predicted pollen limitation from a wider sample of plants with a simpler model than did population size. Specifically, the largest site, a nature preserve, is composed of plants with intermediate individual isolation and, as predicted, intermediate pollen limitation, but its large population size poorly predicted population mean pollen limitation.     

Expanding the limits of the pollen-limitation concept: effects of pollen quantity and quality

Pollination commonly limits seed production, as addition of pollen to stigmas often increases fecundity. This response is usually interpreted as evidence that plants' stigmas receive too few pollen grains to maximize ovule fertilization (quantity limitation); however, many genetic studies demonstrate that poor-quality pollen can also reduce seed production (quality limitation). We explore both aspects of pollen limitation theoretically with a dose-response model that incorporates a saturating negative-exponential relation of seed production to pollen receipt. This relation depends on aspects of ovule production, pollen import, pollen-pistil interactions and seed development, all of which can contribute to pollen limitation. Our model reveals that quantity limitation is restricted to the lowest range of pollen receipt, for which siring success per pollen grain is high, whereas quality limitation acts throughout the range of pollen receipt if plants do not import the highest-quality pollen. In addition to pollinator availability and efficiency, quantity limitation is governed by all post-pollination aspects of seed production. In contrast, quality limitation depends on the difference in survival of embryos sired by naturally delivered pollen vs. by pollen of maximal quality. We briefly illustrate the distinction between these two components of pollen limitation with results from the mistletoe Tristerix corymbosus. Our model also shows that the standard pollen-supplementation technique neither estimates the total intensity of pollen limitation nor distinguishes between its quantity and quality components. As an alternative, we propose a methodological protocol that requires both measurement of seed production following excess pollination with only outcross pollen and quantification of the dose-response relation of seed output to pollen receipt. This method estimates both the total extent of pollen limitation and its two components. Finally, we consider the influences on quantity and quality limitation, which reveals that quantity limitation probably occurs much less often than has been inferred from pollen-supplementation experiments. These interpretations suggest that an expanded perspective that recognizes the fecundity consequences of pollination with poor-quality pollen would promote ecological understanding of pollen limitation.     

Recruitment limitation constrains local species richness and productivity in dry grassland

Species coexistence and local-scale species richness are limited by the availability of seeds and microsites for germination and establishment. We conducted a seed addition experiment in seminatural grassland at three sites in southern Switzerland and repeated the experiment in two successive years to evaluate various circumstances under which seed limitation and establishment success affect community functioning. A collection of 144,000 seeds of 22 meadow species including grasses and forbs of local provenance was gathered, and seeds were individually sown in a density that resembled natural seed rain. The three communities were seed limited. Three years after sowing, single species varied in emergence (0-50%), survival (0-69%), and establishment rates (0-27%). One annual and 13 perennial species reached reproductive stage. Low establishment at one site and reduced growth at another site indicated stronger microsite limitation compared to the third site. Recruitment was influenced by differences in abiotic environmental conditions between sites (water availability, soil minerals) and by within-site differences in biotic interaction (competition). At the least water-limited site, sowing resulted in an increase in phytomass due to establishment of short-lived perennials in the second and third years after sowing. This increase persisted over the following two years due to establishment of longer-lived perennials. After sowing in a wetter year with higher phytomass, however, productivity did not increase, because higher intensity of competition in an early phase of establishment resulted in less vigorous plants later on. Due to the generally favorable weather conditions during this study, sowing year had a small effect on numbers of established individuals over all species. Recruitment limitation can thus constrain local-scale species richness and productivity, either by a lack of seeds or by reduced seedling growth, likely due to competition from the established vegetation.     

Tracing impacts of partner abundance in facultative pollination mutualisms: from individuals to populations

Partner abundance affects costs and benefits in obligate mutualisms, but its role in facultative partnerships is less clear. We address this gap in a pollination web consisting of two clovers (Trifolium) that differ in specialization on a bumble bee pollinator Bombus balteatus. We examine how pollination niche breadth affects plant responses to pollinator abundance, comparing early-flowering (specialized) and late-flowering (generalized) cohorts of T. parryi and early T. parryi to T. dasyphyllum, a pollination generalist. Co-pollinators disrupt the link between B. halteatus visitation and pollination rate for both clovers. Only for early-flowering T. parryi do visitation, pollination, and seed set increase with density of B. balteatus. Bumble bee density also alters timing of seed germination in T. parryi, with seeds from plants receiving augmented B. balteatus germinating sooner than seeds of open-pollinated counterparts. Benefits saturate at intermediate bumble bee densities. Despite strong effects of B. balteatus density on individual plant fitness components, population models suggest little impact of B. balteatus density on lamda in T. parryi or T. dasyphyllum. Findings show that functional redundancy in a pollinator guild mediates host-plant responses to partner density. Unexpected effects of pollinator density on life history schedule have implications for recruitment under pollinator decline.     

Effects of Habitat Size and Patch Isolation on Reproductive Success of the Serpentine Morning Glory

Abstract: We examined the effects of habitat area and patch isolation on reproductive success in serpentine morning glory ( Calystegia collina [Convolvulaceae]), a primarily self-incompatible clonal plant endemic to serpentine outcrops in northern California's coast ranges. Within a 4000-km ² region, we compared the reproductive success of C. collina on 16 small (<5 ha) and 7 large ( >300 ha) outcrops. Flower and fruit production were significantly higher on large serpentine outcrops than on small outcrops. Fruit production also was positively correlated with the soil's ratio of calcium to magnesium. Successful pollination was positively affected by flower density and the number of other flowering patches within 100 m of a C. collina patch. The number of nearby flowering patches was considerably higher on large than on small outcrops. Flowers on large outcrops did not receive significantly more bee visitors than flowers on small outcrops, suggesting that pollination success is related to the quality rather than the quantity of pollen deposited. Fruit production by plants on both small and large outcrops was enhanced by the experimental addition of pollen from other patches, but not by the addition of pollen from the same patch. These findings demonstrate that the size of habitat may have strong effects on the reproductive success of locally endemic plants by enhancing opportunities for successful sexual reproduction. They also warn against the presumption that naturally patchy plant species are invulnerable to the effects of habitat fragmentation.     

番茄某些繁殖特性对增强UV-B辐射的响应

研究了增强UV-B辐射对大田生长的两种番茄(早熟型“同辉”和晚熟型“霞光”)某些繁殖特性(最大开花数、花粉萌发和花粉管伸长、果实产量、果实品质以及子代种子的数量和质量)的影响.结果表明,与对照相比,“同辉”开花数在高辐射下增加,“霞光”开花数在低辐射下减少;“同辉”果实产量在两种辐射下都明显增加,而“霞光”果实产量只在低辐射下增加;辐射抑制了“同辉”的花粉萌发和花粉管伸长,而只是降低了“霞光”的花粉萌发率;强辐射造成两种番茄果实番茄红素下降。经过一季的增强UV-B辐射处理,“同辉”种子数量增加,低辐射下种子明显变小;“霞光”种子数量减少,且明显变小.就开花数、果实产量和种子质量而言,早熟的“同辉”品种更具抗性,而晚熟的“霞光”更敏感.图3表1参29     

Pollinator effectiveness varies with experimental shifts in flowering time

The earlier flowering times exhibited by many plant species are a conspicuous sign of climate change. Altered phenologies have caused concern that species could suffer population declines if they flower at times when effective pollinators are unavailable. For two perennial wildflowers, Tradescantia ohiensis and Asclepias incarnata, we used an experimental approach to explore how changing phenology affects the taxonomic composition of the pollinator assemblage and the effectiveness of individual pollinator taxa. After finding in the previous year that fruit set varied with flowering time, we manipulated flowering onset in greenhouses, placed plants in the field over the span of five weeks, and measured pollinator effectiveness as the number of seeds produced after a single visit to a flower. The average effectiveness of pollinators and the expected rates of pollination success were lower for plants of both species flowering earlier than for plants flowering at historical times, suggesting there could be reproductive costs to earlier flowering. Whereas for A. incarnata, differences in average seed set among weeks were due primarily to changes in the composition of the pollinator assemblage, the differences for T. ohiensis were driven by the combined effects of compositional changes and increases over time in the effectiveness of some pollinator taxa. Both species face the possibility of temporal mismatch between the availability of the most effective pollinators and the onset of flowering, and changes in the effectiveness of individual pollinator taxa through time may add an unexpected element to the reproductive consequences of such mismatches.     

Linkages between phenology, pollination, photosynthesis, and reproduction in deciduous forest understory plants

Light availability in the understory of deciduous forests changes drastically within the growing season due to the foliage dynamics of canopy trees. Because flowering phenology, photosynthetic characteristics, and fruiting success respond to such strong seasonality in light availability, we hypothesized that understory plants in such ecosystems should describe distinct phenological groups or syndromes where "syndrome" is defined only as a set of characteristics that co-occur. To identify these phenological syndromes, we studied the flowering phenology, fruit or seed set, and photosynthetic characteristics for 18 perennial understory herbaceous species that differed in reproductive strategy over eight years in a deciduous forest in northern Japan. Three phenological groups emerged from this study: (1) spring bloomers, flowering and fruiting before the completion of canopy closure; (2) early-summer bloomers, flowering during the progress of canopy closure and fruiting after canopy closure; and (3) late-summer bloomers, flowering and fruiting after canopy closure. The spring bloomers had high photosynthetic rates and high fruiting abilities, but the flowering time varied considerably among years due to yearly fluctuations of snowmelt date. Bumble bee-pollinated species of spring bloomers showed variable seed-set success, while fly-pollinated species showed relatively stable seed sets over the years. The early-summer bloomers showed low fruiting abilities irrespective of pollination success, reflecting severe resource limitation with decelerating light availability during fruit development. Although the late-summer bloomers showed low photosynthetic rates under low-light conditions, high fruit-set success was attained if pollination was sufficient. These results support our hypothesis that phenological syndromes may be found in deciduous forest understory plants. Given that reproductive success of bee-pollinated spring bloomers is highly susceptible to seasonal fluctuation, climate change may have its strongest impacts on this group.     

Anther-mimicking floral guides exploit a conflict between innate preference and learning in bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>)

Bee-pollinated plants are frequently dichogamous: e.g. each flower has a discernable male and female phase, with only the male phase offering a pollen reward. Pollen-collecting bees should therefore discriminate against female-phase flowers to maximise their rate of pollen harvest, but this behaviour would reduce plant fitness due to inferior pollination. Here, we test the hypothesis that flowers use pollen-mimicking floral guides to prevent flower-phase discrimination. Such floral guides resemble pollen in spectral reflection properties and are widespread among angiosperm flowers. In an array of artificial flowers, bumblebees learned less well to discriminate between flower variants simulating different flowering phases when both flower variants carried an additional pollen-yellow guide mark. This effect depended crucially on the pollen-yellow colour of the guide mark and on its spatial position within the artificial flower. We suggest that floral guides evolved to inhibit flower-phase learning in bees by exploiting the innate colour preferences of their pollinators.     

Potential Misuse of Avian Density as a Conservation Metric

Abstract: Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual‐ and population‐level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density‐independent and density‐dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density‐dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.     

Effects of Population Size on Seed Production and Germinability in an Endangered, Fragmented Grassland Plant

Abstract: Fragmentation and isolation of plant populations are thought to affect demographic processes such as seed production and cause reductions in fitness. I followed seed set over a 3-year period in eight populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) that differed in population size from 13 to over 5000 flowering plants. Germinability of the resultant seed was also examined to determine whether small populations had lower fitness than large populations. Seed set was significantly associated with population size in 2 of the 3 years. Small populations (<30 flowering plants) produced significantly fewer seeds per head in 1994 and 1995 than did large populations (500 to over 5000 flowering plants), which did not differ significantly from one another. There was, however, substantial variation within populations. In 1993 seed production did not follow any simple relationship with population size, possibly because environmental stress from low rainfall had an overriding impact. Differences in seed germinability between populations were largely not evident, suggesting that this aspect of fitness has not declined substantially in small populations relative to large populations. This study suggests that nongenetic, demographic factors are of immediate importance to the persistence of small populations of R. leptorrhynchoides because of their potential impacts on seedling recruitment.     

Postdispersal seed predation limits the abundance of a long-lived perennial forb (Lithospermum ruderale)

Loss of seeds to consumers is common in plant communities, but the degree to which these losses influence plant abundance or population growth is often unclear. This is particularly the case for postdispersal seed predation by rodents, as most studies of rodent seed predation have focused on the sources of spatiotemporal variation in seed loss but not quantified the population consequences of this loss. In previous work we showed that seed predation by deer mice (Peromyscus maniculatus) substantially reduced seedling recruitment and establishment of Lithospermum ruderale (Boraginaceae), a long-lived perennial forb. To shed light on how rodent seed predation and the near-term effects on plant recruitment might influence longer-term patterns of L. ruderale population growth, we combined experimental results with demographic data in stage-based population models. Model outputs revealed that rodent seed predation had a significant impact on L. ruderale population growth rate (lambda). With the removal of postdispersal seed predation, the projected population growth rates increased between 0.06 and 0.12, depending on site (mean deltalambda across sites = 0.08). Seed predation shifted the projected stable stage distribution of populations from one with a high proportion of young plants to one in which larger adult size classes dominate. Elasticities of vital rates also changed, with germination and growth of seedlings and young plants becoming more important with the removal of seed predation. Simulations varying the magnitude of seed predation pressure while holding other vital rates constant showed that seed predation could lower lambda even if only 40% of available seeds were consumed. These results demonstrate that rodent granivory can be a potent force limiting the abundance of a long-lived perennial forb.     

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.     

Effect of Forest Fragmentation on Genetic Diversity and Mating System in a Tropical Tree, Pithecellobium elegans

Genetic diversity, population differentiation, and temporal variation in outcrossing rates were examined for Pithecellobium elegans , a Neotropical rain forest canopy tree. Several forest fragments and a large reserve (1500 ha) were compared for several population genetic parameters. For eight populations sampled on the Atlantic coastal plain of Costa Rica, allozyme heterozygosity (0.13), polymorphism (35%), and effective number of alleles (1.24) were similar to values reported for other tropical tree species that occur at similar densities of less than one individual per hectare. These measures of genetic variation were lowest in populations of the smallest size, farthest from the reserve, and more isolated from other populations. Differentiation among samples collected in small forest fragments and the reserve population accounted for 10% of the total genetic variation observed. There was a positive relationship between the level of differentiation of populations from the reserve population and their distance from the reserve. Though predominantly an annually flowering species, the number of trees in flower at any one time varied from 80% of observed trees to only 6%. Outcrossing rates did not differ for two episodes in which the proportions of flowering trees were 33% and 80%. But periods of low density of flowering adults resulted in poor seed crops or failure to set fruit for many individuals. Population size at many sites will be effectively decreased because of the variation in flowering. Fragmentation of what was once a large, continuous forested area is resulting in genetic erosion of small, isolated populations of Pithecellobium elegans .     

Imperfect Replacement of Native Species by Non‐Native Species as Pollinators of Endemic Hawaiian Plants

Native plant species that have lost their mutualist partners may require non‐native pollinators or seed dispersers to maintain reproduction. When natives are highly specialized, however, it appears doubtful that introduced generalists will partner effectively with them. We used visitation observations and pollination treatments (experimental manipulations of pollen transfer) to examine relationships between the introduced, generalist Japanese White‐eye (Zosterops japonicus) and 3 endemic Hawaiian plant species (Clermontia parviflora, C. montis‐loa, and C. hawaiiensis). These plants are characterized by curved, tubular flowers, apparently adapted for pollination by curve‐billed Hawaiian honeycreepers. Z. japonicus were responsible for over 80% of visits to flowers of the small‐flowered C. parviflora and the midsize‐flowered C. montis‐loa. Z. japonicus‐visited flowers set significantly more seed than did bagged flowers. Z. japonicus also demonstrated the potential to act as an occasional Clermontia seed disperser, although ground‐based frugivory by non‐native mammals likely dominates seed dispersal. The large‐flowered C. hawaiiensis received no visitation by any birds during observations. Unmanipulated and bagged C. hawaiiensis flowers set similar numbers of seeds. Direct examination of Z. japonicus and Clermontia morphologies suggests a mismatch between Z. japonicus bill morphology and C. hawaiiensis flower morphology. In combination, our results suggest that Z. japonicus has established an effective pollination relationship with C. parviflora and C. montis‐loa and that the large flowers of C. hawaiiensis preclude effective visitation by Z. japonicus. Remplazo Imperfecto de Especies Nativas por Especies No‐Nativas como Polinizadores de Plantas Endémicas de Hawaii     

Meta-analysis of susceptibility of woody plants to loss of genetic diversity through habitat fragmentation

Shrubs and trees are assumed less likely to lose genetic variation in response to habitat fragmentation because they have certain life-history characteristics such as long lifespans and extensive pollen flow. To test this assumption, we conducted a meta-analysis with data on 97 woody plant species derived from 98 studies of habitat fragmentation. We measured the weighted response of four different measures of population-level genetic diversity to habitat fragmentation with Hedge's d and Spearman rank correlation. We tested whether the genetic response to habitat fragmentation was mediated by life-history traits (longevity, pollination mode, and seed dispersal vector) and study characteristics (genetic marker and plant material used). For both tests of effect size habitat fragmentation was associated with a substantial decrease in expected heterozygosity, number of alleles, and percentage of polymorphic loci, whereas the population inbreeding coefficient was not associated with these measures. The largest proportion of variation among effect sizes was explained by pollination mechanism and by the age of the tissue (progeny or adult) that was genotyped. Our primary finding was that wind-pollinated trees and shrubs appeared to be as likely to lose genetic variation as insect-pollinated species, indicating that severe habitat fragmentation may lead to pollen limitation and limited gene flow. In comparison with results of previous meta-analyses on mainly herbaceous species, we found trees and shrubs were as likely to have negative genetic responses to habitat fragmentation as herbaceous species. We also found that the genetic variation in offspring was generally less than that of adult trees, which is evidence of a genetic extinction debt and probably reflects the genetic diversity of the historical, less-fragmented landscape.