Lattice population dynamics for plants with dispersing seeds and Vegetative propagation

The population dynamics of plants in a lattice structured habitat are studied theoretically. Plants are assumed to propagate both by producing seeds that scatter over the population and by vegetative reproduction by extending runners, rhizomes, or roots, to neighboring vacant sites. In addtion, the seed production rate may be dependent on the local density in the neighborhood, indicating beneficial or harmful crowding effects. Two sets of population dynamical equation(s) are derived: one based onmean-field approximation and the other based onpair approximation (tracing both global and local densities simultaneously). We examine the accuracy of these approximate dynamics by comparing them with direct computer simulation of the stochastic lattice model. Pair approximation is much more accurate than mean-field approximation. Mean-field approximation overestimates the parameter range for persistence if crowding effects on seed production are harmful or weakly beneficial, but underestimates it if crowding effects are highly beneficial. Dynamics may show bistability (both population persistence and extinction) if the effect of crowding is strongly beneficial. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximised by a mixture of seed production and vegetative reproduction, rather than by pure seed production or by pure vegetative reproduction. This result is correctly predicted by pair approximation but not by mean-field approximation.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

生境破碎对古尔班通古特沙漠南缘骆驼刺种群生物学特性的影响

在破碎化生境中,植物种群的生物学特性直接反映植物对环境的响应特征和适应对策,探讨不同破碎化程度生境中植株的大小、密度、结实率、种子大小、种子质量以及萌发特性等,对深入了解植物在不同生境中的生存和繁殖状况有重要的指导意义。对古尔班通古特沙漠南缘9个斑块化骆驼刺种群的形态特征和繁殖生物学参数进行了连续2年的调查研究,研究结果表明：随着生境破碎化程度的加剧,骆驼刺的株高、冠幅、密度、结实率、种子大小、种子质量以及萌发率等均显著下降（P < 0.05）。生境破碎不仅强烈抑制了骆驼刺植株的营养生长,使其生长矮小,个体分布稀疏,而且严重阻碍了骆驼刺的繁殖成功和种群更新,增加了种群衰退甚至灭绝的风险。     

Fitness and survival in fragmented populations of Narthecium ossifragum at the species' range margin

Decreasing population size and increasing isolation may lead to reduced plant fitness. These effects of habitat fragmentation may especially apply to populations close to the margin of their geographical range, where populations generally are often smaller and more isolated, and the environment is less favourable than at the range centre. In this context we studied north-west German range-margin populations of Narthecium ossifragum (L.) HUDS., a clonal, perennial herb typical for acidic and nutrient-poor wet heathlands. We asked whether plant fitness and reproduction of the range-margin populations were affected by population size and habitat quality, and whether any changes in population size over the past 10–20 years were related to past population size and habitat quality.Population size varied between 60 and 100.000 individuals and was highly positively related to shoot density. Larger (and denser) populations formed more flowers and fruits, but fewer seeds. Soil water content had a strong positive effect on seed production and seed mass, and the latter also increased with increasing soil phosphorus content. Relative light intensity showed a positive impact on shoot density and, hence, population size. Compared to British range centre populations, the populations at the German range margin formed more, but smaller seeds that failed to germinate both in the field and in laboratory experiments. Despite the lack of sexual reproduction, population sizes generally had not decreased during the past two decades, most likely due to successful clonal growth. The change in population size of Narthecium, however, was negatively related to the present soil phosphorus content, indicating that an increased supply of nutrients and a decrease in light availability may have a negative effect on population dynamics.The results imply that fragmented and small range-margin populations not necessarily are doomed to extinction, provided that the habitat quality remains suitable for growth and vegetative reproduction.     

Predation and infanticide influence ideal free choice by a parrot occupying heterogeneous tropical habitats

The ideal free distribution (IFD) predicts that organisms will disperse to sites that maximize their fitness based on availability of resources. Habitat heterogeneity underlies resource variation and influences spatial variation in demography and the distribution of populations. We relate nest site productivity at multiple scales measured over a decade to habitat quality in a box-nesting population of Forpus passerinus (green-rumped parrotlets) in Venezuela to examine critical IFD assumptions. Variation in reproductive success at the local population and neighborhood scales had a much larger influence on productivity (fledglings per nest box per year) than nest site or female identity. Habitat features were reliable cues of nest site quality. Nest sites with less vegetative cover produced greater numbers of fledglings than sites with more cover. However, there was also a competitive cost to nesting in high-quality, low-vegetative cover nest boxes, as these sites experienced the most infanticide events. In the lowland local population, water depth and cover surrounding nest sites were related with F. passerinus productivity. Low vegetative cover and deeper water were associated with lower predation rates, suggesting that predation could be a primary factor driving habitat selection patterns. Parrotlets also demonstrated directional dispersal. Pairs that changed nest sites were more likely to disperse from poor-quality nest sites to high-quality nest sites rather than vice versa, and juveniles were more likely to disperse to, or remain in, the more productive of the two local populations. Parrotlets exhibited three characteristics fundamental to the IFD: habitat heterogeneity within and between local populations, reliable habitat cues to productivity, and active dispersal to sites of higher fitness.     

扩散对破碎化景观上宿主-寄生种群动态的影响

景观破碎化和扩散是空间种群模型的重要因素,对生物入侵存在着深远的影响。本章将基于偶对近似模型,探讨由局部和全局宿主-寄生相互作用共同决定的扩散模式对破坏性景观上疾病入侵与传播的影响。其中,生境破坏由生境丧失量与生境破碎化程度来描述。模拟结果显示,宿主和病毒的全局扩散对疾病的入侵与种群密度产生不对称效应：病毒的全局扩散对系统产生的影响较宿主的全局扩散更为显著。不同扩散模式下,生境丧失越高或破碎化程度越低,均将越有害于寄生病毒的入侵;同时,生境的破坏程度也显著地影响了入侵阈值对扩散模式的响应机制。本文研究结果暗示,景观破碎化的空间分布格局以及病毒扩散的限制均可作为物种保护与管理中有效的疾病控制策略。该研究结果在一定意义上丰富和发展了寄生感染理论,为物种保护提供了生态学理论依据。     

Negative effects of habitat degradation and fragmentation on the declining grassland plant Trifolium montanum

Changes in land use have resulted in a strong decline in the plant diversity of nutrient-poor grasslands, but little is known about the combined effects of habitat degradation and fragmentation on populations of individual species. We studied these effects on stage structure, recruitment, reproduction and offspring fitness in populations of the declining perennial grassland plant Trifolium montanum in central Germany. Density and survival probability of juvenile plants decreased with light competition, measured as leaf area index (LAI) above T. montanum plants, resulting in aged populations with few juvenile plants at unmanaged sites with higher LAI. Reproduction of T. montanum was not related to LAI, but increased strongly with local density, suggesting pollinator limitation in fragmented populations with a low density of flowering plants. In the common garden, the survival of sown offspring increased with mean seed size, whereas seed production of offspring decreased with isolation, and in strong contrast to previous studies, also decreased with size and density of the population of origin. This could be due to increased inbreeding because of pollination between closely related neighbouring plants in dense and large populations. Our results indicate that both habitat degradation and fragmentation have negative effects on populations of T. montanum, but affect different phases of the life cycle. In the short term, the effects of habitat degradation are more important than those of fragmentation, and populations of T. montanum are primarily threatened by an increase in light competition in unmanaged sites, which rapidly affects the dynamics of the populations. The observed opposite effects of habitat fragmentation on reproduction and offspring fitness indicate that the effects of population size, density and isolation on plant fitness and population viability may be complex.     

Linking vital rates to invasiveness of a perennial herb

Invaders generally show better individual performance than non-invaders and, therefore, vital rates (survival, growth, fecundity) could potentially be used to predict species invasiveness outside their native range. Comparative studies have usually correlated vital rates with the invasiveness status of species, while few studies have investigated them in relation to population growth rate. Here, I examined the influence of five vital rates (plant establishment, survival, growth, flowering probability, seed production) and their variability (across geographic regions, habitat types, population sizes and population densities) on population growth rate (λ) using data from 37 populations of an invasive, iteroparous herb (Lupinus polyphyllus) in a part of its invaded range in Finland. Variation in vital rates was often related to habitat type and population density. The performance of the populations varied from declining to rapidly increasing independently of habitat type, population size or population density, but differed between regions. The population growth rate increased linearly with plant establishment, and with the survival and growth of vegetative individuals, while the survival of flowering individuals and annual seed production were not related to λ. The vital rates responsible for rapid population growth varied among populations. These findings highlight the importance of both regional and local conditions to plant population dynamics, demonstrating that individual vital rates do not necessarily correlate with λ. Therefore, to understand the role of individual vital rates in a species ability to invade, it is necessary to quantify their effect on population growth rate.     

Variations in sexual and asexual reproduction of Scirpus mariqueter along an elevational gradient

In order to assess the importance of sexual and asexual reproduction during the life history of Scirpus mariqueter, its reproductive and growth characters were concurrently examined along an elevational gradient (from low elevation to high elevation). The proportions of flowering shoot and inflorescence mass, seed : flower ratio and seed weight were used to quantify the investment in sexual reproduction. The proportions of current-year shoot and rhizome mass were used to quantify the investment in asexual reproduction, and the proportion of corm mass was used for growth, respectively. It was found that vegetative propagation predominated at low elevation, whereas sexual reproduction predominated at high elevation; and that sexual reproduction increased with declining asexual reproduction along the gradient. The results suggest that asexual reproduction is relatively favored in the early life stage, whereas sexual reproduction is favored when the population becomes mature and aged, probably because of the functional differentiation between the two reproductive types. Sexual productive characters (i.e. the proportions of flowering shoot and inflorescence mass) were negatively correlated to both growth and asexual reproductive characters along the gradient, indicating there might exist some trade-offs among growth, sexual and asexual reproduction during the life history. However, no obvious pattern was found between asexual reproductive characters and growth characters along the elevational gradient, possibly because of the varied relationships between them at different life stages. The variations in sexual and asexual reproduction in the species and the relationship between them are thought to be of great significance for local population growth, species persistence and evolution.     

Habitat and microsite influence demography of two herbs in intact and degraded scrub

Identifying environmental factors associated with vital rate variation is critical to predict population consequences of environmental perturbation. We used matrix models to explore effects of habitat and microsite on demography of two widespread herbs, Chamaecrista fasciculata (partridge pea) and Balduina angustifolia (yellow buttons). We evaluated models simulating population dynamics in common microsites (shrub, litter, bare sand) within two habitats (intact, degraded Florida scrub) using data on experimental populations initiated by sowing seeds, and natural seed production. Models included four stages (seed bank, small vegetative, large vegetative, reproductive) and three vital rates (survival, growth, fecundity), summarized in sixteen transitions. We conducted life table response experiments to assess contributions of each habitat and microsite to population growth rates. We found that (1) C. fasciculata had greatest population growth in degraded habitat and litter microsites, (2) B. angustifolia had similar population growth between habitats and greatest in bare sand microsites, (3) advancing growth transitions of C. fasciculata had greatest elasticity on population growth in degraded habitat, shrub, and litter, as did seed survival in intact habitat and bare sand, (4) seed survival and advancing growth transitions of B. angustifolia had greatest elasticity on population growth in both habitats, as did seed survival in shrub and litter, and advancing growth in bare sand. Greater population growth of C. fasciculata in degraded scrub is probably explained by release from belowground competition; B. angustifolia may be most affected by competition with shrubs. Microsites in intact scrub were not ecologically equivalent to those in degraded scrub emphasizing that intact scrub is ecologically complex and critical to preserve.     

Warm temperature conditions restrict the sexual reproduction and vegetative growth of the spring ephemeral <Emphasis Type="Italic">Gagea lutea</Emphasis> (Liliaceae)

The responses of reproduction and growth to climate warming are important issues to predict the fate of plant populations at high latitudes. Spring ephemerals inhabiting cool-temperate forests grow better under cool conditions, but how reproductive performance is influenced by warm weather is unclear. The phenological and physiological responses of reproduction and vegetative growth to warm temperature and light conditions were evaluated in the spring ephemeral Gagea lutea. Leaf and bract physiological activities, bulb growth, and seed production were compared among reproductive plants grown in forest, open, and greenhouse (GH; warming manipulation in the open site) plots. In vitro pollen germination ability was tested under various temperatures. In the GH, leaf and bract photosynthetic activities decreased rapidly at the fruiting stage, but dark respiration rates remained high, resulting in higher carbon exhaust in warm conditions. Both leaf and bract sizes and their longevities were reduced in the GH. Annual bulb growth was largest in the forest plot and smallest in the GH plot. Pollen germination was strongly inhibited at high temperature (30 °C). Fruit and seed productions were decreased only in the GH plot. Both vegetative and reproductive activities were negatively affected by warm temperature, resulting in less vegetative growth and lower seed-set, whereas an understory habitat was beneficial for vegetative growth and showed similar seed production to an open habitat over the experimental period. Decreasing population dynamics of spring ephemerals was predicted in response to future warming climate not only by growth inhibition but also by restriction of seed production.     

Bud banks of perennial savanna grasses in Botswana

Three semi‐arid savanna grasses in Botswana (Stipagrostis uniplumis, Eragrostis lehmanniana, and Aristida stipitata) were sampled to quantify their belowground bud banks during the dormant season and to estimate their relative allocation to vegetative and sexual reproduction. Bud banks of these African perennial caespitose grasses were also compared with four perennial caespitose grasses of semi‐arid North American grasslands. The three African grasses each maintained approximately two buds per tiller and showed a high percentage (88–99%) of tillers producing seed. Only E. lehmanniana produced new aerial tillers from axillary buds at elevated nodes on the stem as well as from the belowground bud bank. Compared with species of North American grasslands, these African grasses produced fewer belowground buds but showed a much higher percentage of tillers producing seed. These patterns indicate relatively greater belowground meristem limitation, lower allocation to vegetative reproduction (tillering) and higher allocation to seed reproduction in these African grasses, although studies of more species are needed to assess the generality of this pattern. The management of savannas in ways that favour the maintenance of a reserve population of belowground buds may increase the ability of grasses to respond to pulses of resource availability, increase their compensatory growth capacity following grazing or drought, and decrease the invasibility of these plant communities by exotic species, whereas maintaining allocation to sexual reproduction may be important for conserving genetic variation and enhancing their capacity to adapt to environmental change.     

Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae)

Syneilesis palmata reproduces by both seeds and vegetative propagules (short rhizomes). The latter result in the production of new plants that are larger in size and hence have a higher survival probability and a higher growth rate than seeds. A previous study predicted that the optimal reproductive strategy, in terms of maximizing population growth rate (a fitness measure under no density regulations), was pure vegetative reproduction. However, high resource investment to vegetative propagules can cause local crowding resulting in reduced demographic performances of the plants, because the vegetative propagules of Syneilesis are produced close to one another. We examined, in this situation, the impact of allocating a certain proportion of reproductive resource to seeds with relatively greater capacity for dispersal. We simulated dynamics of hypothetical Syneilesis populations with various reproductive resource allocation balances (from pure seed to pure vegetative reproduction), using a density-dependent matrix model. In the model, it was assumed that plants from vegetative propagules experienced density-dependent reduction in their survival probabilities, but this was not the case for plants originating from seeds. Each allocation strategy was evaluated based on an equilibrium population density, a fitness measure under density-dependent regulations. The optimal reproductive strategy predicted was pure vegetative reproduction. Unrealistic conditions were required for seed reproduction to be favoured, such as the production of seeds one hundred times the normal number per unit resource investment. However, the conditions were fairly relaxed compared with those required in the model where no density effects were incorporated. This indicates that escape from local crowding is likely to be one of the roles of seed production in Syneilesis.     

Change in spatial distribution patterns of a biennial plant between growth stages and generations in a patchy habitat

BACKGROUND AND AIMS: The aim of the study was to evaluate factors causing change in spatial distribution patterns of plants between growth stages and generations for a monocarpic biennial plant, Lysimachia rubida. It was assumed that habitat heterogeneity was a primary factor determining spatial patterns of plants, and a randomization procedure was developed for testing the null hypothesis that only spatial association with ground surface conditions determined spatial patterns of plants. METHODS: A 5-year demographic census was conducted on an open dry habitat that was heterogeneous with regard to the ground surface conditions. KEY RESULTS: There was significant habitat association in that plants at vegetative and reproductive stages were denser in areas with smaller gravel than with larger gravel. Point process analyses rejected the null hypothesis of the spatial association with ground surface conditions. CONCLUSIONS: The results suggest that other factors, such as patchy seed dispersal, secondary dispersal of the seeds and life-history variation at various spatial scales, also affected spatial patterns of individuals in a population of L. rubida. Spatial structures and dynamics of a local population in a patchy habitat represent various performances of plants within patches and seed dispersal within a patch and beyond the patch.     

Limits to local spatial spread in a highly invasive annual grass (Microstegium vimineum)

The potential roles of seed and microsite limitation in local spatial spread of the invasive grass Microstegium vimineum were experimentally investigated in a woodland and open lawn in central New Jersey, USA. Plots (30 × 30 cm) in three sites previously unoccupied by M. vimineum for at least 8 years (woodland interior, woodland edge, and open lawn) were sown with ~262 M. vimineum seeds in early spring 2008. Seedling emergence, density, summer growth and autumn reproduction were compared to plots in a nearby control population where natural recruitment occurred. Seedling emergence was greatest in the open lawn (54% of seeds sown) where plants showed the greatest growth and reproduction due to high light availability. Seedling emergence was lowest in the woodland interior (24%) and edge sites (9%), and growth and reproduction were greatly reduced there (relative to the control). Plots in the open lawn supported a consistently high density of M. vimineum (>1,000 plants per m²) through the growing period (April to October). The number of seeds in both cleistogamous and chasmogamous spikelets was correlated with shoot dry mass and thus, total seed production was greatest in the sunny open lawn where plants were largest, despite high density there. Across all sites, plants in plots at the highest densities produced the most seeds. Total seed production correlated with levels of light, but not soil moisture. Both seed availability and microsite limitation may reduce the probability of establishment of new M. vimineum populations into previously unoccupied sites. Intraspecific density does not negatively affect survival or reproduction. Light and soil moisture can be limiting abiotic factors in some areas, but poor, natural seed dispersal limits the distribution of this invasive species on a local scale.     

Local forest environment largely affects below-ground growth, clonal diversity and fine-scale spatial genetic structure in the temperate deciduous forest herb Paris quadrifolia

Paris quadrifolia (herb Paris) is a long-lived, clonal woodland herb that shows strong differences in local population size and shoot density along an environmental gradient of soil and light conditions. This environmentally based structuring may be mediated by differences in clonal growth and seedling recruitment through sexual reproduction. To study the interrelationship between environmental conditions and spatial patterns of clonal growth, the spatial genetic structure of four P. quadrifolia populations growing in strongly contrasting sites was determined. In the first place, plant excavations were performed in order to (i) determine differences in below-ground growth of genets, (ii) investigate connectedness of ramets and (iii) determine total genet size. Although no differences in internode length were found among sites, clones in moist sites were much smaller (genets usually consisted of 1-3 interconnected shoots, most of them flowering) than genets in dry sites, which consisted of up to 15 interconnected shoots, the majority of which were vegetative. Further, amplified fragment length polymorphism (AFLP) markers were used. Clonal diversity was higher in populations located in moist and productive ash-poplar forests compared to those found in drier and less productive mixed forest sites (G/N: 0.27 and 0.14 and Simpson's D: 0.84 and 0.75, respectively). Patterns of spatial population genetic structure under dry conditions revealed several large clones dominating the entire population, whereas in moist sites many small genets were observed. Nevertheless, strong spatial genetic structure of the genet population was observed. Our results clearly demonstrate that patterns of clonal diversity and growth form of P. quadrifolia differ among environments. Limited seedling recruitment and large clone sizes due to higher connectedness of ramets explain the low clonal diversity in dry sites. In moist sites, higher levels of clonal diversity and small clone sizes indicate repeated seedling recruitment, whereas strong spatial genetic structure suggests limited seed dispersal within populations.     

The role of vegetative spread and seed dispersal for optimal life histories of clonal plants: a simulation study

Sexual and vegetative reproduction of clonal plants phenotypically differ in dispersal distance, in the phenology of offspring production and establishment, and in the success of establishment. We applied a combination of analytical and of spatially explicit individual-based simulation modelling to calculate long-term fitness. We predicted optimal clonal plant life histories in a parameter space spanned by stolon number, stolon internode length, and relative allocation to sexual and vegetative reproduction. For a given allocation to sexual reproduction and number of stolons, fitness was optimised for rather short internode lengths under small disturbances, and for the longest possible internodes under larger disturbances. A trade-off between length and number of vegetative spacers drew parameters away from their unrestricted optima. Now, intermediate length and number of spacers led to maximum fitness under large disturbances. Simultaneous trade-offs between sexual and vegetative reproduction and between the length and number of spacers could also lead to fitness optima at intermediate parameter values, depending on the success of seedling establishment. We demonstrated that spatial habitat structure (1) selects for an efficient use of available space either by optimum internode length or by investment into seeds, which disperse farther than vegetative spacers, and (2) leads to an interaction between trade-offs. We conclude, that dispersal distance, i.e. a spatial life-history component, and trade-offs must be included in considerations on adaptive evolution of clonal life histories.     

Mechanisms driving the density–area relationship in a saproxylic beetle

Mechanisms underlying density–area relationships (correlations between population density and patch size) have rarely been tested experimentally. It is often assumed that higher density on large patches is evidence that large patches are high quality (i.e. have greater survival and reproduction), but the same pattern could result from disproportionate movement from small to large patches. Movement-based and within-patch processes must be experimentally separated to show that large patches are indeed of higher quality, but few studies have done so. We experimentally tested movement-based and within-patch hypotheses to explain the positive density–area relationship observed for a saproxylic (decayed wood-dependent) beetle, Odontotaenius disjunctus Illiger (Coleoptera: Passalidae). In separate experiments we quantified (1) immigration into and (2) finite growth rate within logs (=patches) that varied in size and density of conspecific beetles. A log was 11.7-fold [95 % confidence interval (CI) 3.4–40.8) and 10.5-fold (95 % CI 2.7–40.9) more likely to contain a new immigrant if it was large or contained a conspecific pair of beetles, respectively. Neither log size nor conspecific density was associated with changes in finite growth rate that would lead to higher density: decreased log size and increased conspecific density reduced finite growth rate in direct proportion to the loss in available resources per mating pair. We conclude that movement behavior rather than habitat quality is responsible for the positive density–area relationship for O. disjunctus. An important implication of our results is that population density is an imperfect indicator of habitat quality.     

Effects of population density, sex morph, and tree size on reproduction in a heterodichogamous maple, Acer mono, in a temperate forest of Japan

The effects of local population density, sex morph [protogynous (PG) or protandrous (PA)], and individual tree size on the demographic processes of seed production were investigated in a heterodichogamous maple, Acer mono Maxim. var. Marmoratum (Nichols.) Hara f. dissectum, in a temperate forest of Japan. As the distance from conspecific reproductive adults increased, the percentage of immature seed fall and empty seeds increased significantly, indicating higher pollination success along with local population density. Although the difference was not distinct, pollination success was affected by the local population density of the reciprocal sex morph rather than that of both sex morphs. The trees at higher local population density sites suffered higher seed mortality due to predation and decay, and tended to produce smaller seeds. Thus, the impacts of local population density operated both positively and negatively on reproduction. As a factor of individual traits, tree size scarcely affected any demographic processes. On the other hand, sex morph did affect pollination success. Trees of PG type had lower immature seed fall than those of PA type, suggesting that the former has higher efficiency of pollen acceptance than the latter. The results on seed demography presented here partly support previous suggestions that heterodichogamous plants exhibit reciprocal cross-pollination and gender specialization as reproductive traits.     

Habitat quality and population density drive occupancy dynamics of snowshoe hare in variegated landscapes

The influence of habitat quality and population density on occupancy dynamics may surpass that of traditional metrics of area and isolation, but often this is not considered explicitly in studies of spatially structured populations. In landscapes that are not easily characterized as binary habitat/non‐habitat (e.g. variegated landscapes), this influence may be even more important and occur at both local and landscape levels. It follows that occupancy dynamics may be driven by disparate processes depending on how extinction or colonization relate to habitat quality and population density. We examined the relative influence of area, structural isolation, habitat quality, local population density, and neighborhood population density (i.e. population density in the landscape around a site) on the probability of extinction and colonization of snowshoe hare Lepus americanus across an expansive forest mosaic landscape (encompassing the northern third of Idaho). Habitat quality and population density were highly influential in determining extinction and colonization, whereas patch area and isolation were much less important. Sites with heavier vegetative cover at the site or landscape‐level were more likely to be colonized and less likely to go extinct, and sites with greater local population density in the previous time step had lower probability of extinction. Sites embedded in high density neighborhoods also were less likely to go extinct, but not more likely to be colonized. We found a significant interaction between local and neighborhood population density on extinction in 1 yr, suggesting that the strength of demographic rescue may vary dependent on local site densities. Our results add to a growing literature showing that factors outside of structural metrics of area and isolation are important drivers of occupancy dynamics. Given the multi‐scaled influence of habitat quality and population density on occupancy dynamics, our work also indicates that research on snowshoe hare must extend beyond simply assessing local factors to understand the spatial dynamics of populations.