A long-term experiment reveals no trade-off between seed persistence and seedling emergence.

Information on seed persistence and seedling emergence from the soil seed bank is critical for understanding species coexistence and predicting community dynamics. However, quantifying seed persistence in the soil is challenging; thus, its association with other life-history traits is poorly known on a broad scale. Using germination phenology for 349 species in a 42-yr experiment, we quantified the persistence-emergence correlations and their associations with intrinsic regeneration traits using Bayesian phylogenetic multilevel models. We showed no trade-off between seed persistence and seedling emergence. Physically dormant seeds were more persistent but exhibited lower emergence than nondormant seeds. Monocarpic species had both higher persistence and emergence than polycarpic species. Seed mass posed a marginal proxy for persistence, while emergence almost doubled from the smallest to the largest seeds. This study challenges the traditional assumption and is the first demonstration of noncorrelation between persistence and emergence, probably owing to the complexity of regenerative strategies. Species with short persistence and low emergence would be the most vulnerable for in situ conservation. Our analyses of this unique, long-term dataset provide a strong incentive for further experimental studies and a rich data resource for future syntheses.

Identifying predictors of translocation success in rare plant species.

The fundamental goal of a rare plant translocation is to create self-sustaining populations with the evolutionary resilience to persist in the long term. Yet, most plant translocation syntheses focus on a few factors influencing short-term benchmarks of success (e.g., survival and reproduction). Short-term benchmarks can be misleading when trying to infer future growth and viability because the factors that promote establishment may differ from those required for long-term persistence. We assembled a large (n = 275) and broadly representative data set of well-documented and monitored (7.9 years on average) at-risk plant translocations to identify the most important site attributes, management techniques, and species' traits for six life-cycle benchmarks and population metrics of translocation success. We used the random forest algorithm to quantify the relative importance of 29 predictor variables for each metric of success. Drivers of translocation outcomes varied across time frames and success metrics. Management techniques had the greatest relative influence on the attainment of life-cycle benchmarks and short-term population trends, whereas site attributes and species' traits were more important for population persistence and long-term trends. Specifically, large founder sizes increased the potential for reproduction and recruitment into the next generation, whereas declining habitat quality and the outplanting of species with low seed production led to increased extinction risks and a reduction in potential reproductive output in the long-term, respectively. We also detected novel interactions between some of the most important drivers, such as an increased probability of next-generation recruitment in species with greater seed production rates, but only when coupled with large founder sizes. Because most significant barriers to plant translocation success can be overcome by improving techniques or resolving site-level issues through early intervention and management, we suggest that by combining long-term monitoring with adaptive management, translocation programs can enhance the prospects of achieving long-term success.

Identificación de pronosticadores del éxito de reubicación en especies raras de plantas Resumen El objetivo fundamental de la reubicación de plantas raras es la creación de poblaciones autosuficientes con resiliencia evolutiva que persistan a la larga. De todas maneras, la mayoría de las síntesis de estas reubicaciones se enfocan en unos cuantos factores que influyen sobre los parámetros a corto plazo del éxito (supervivencia y reproducción). Los parámetros a corto plazo pueden ser engañosos si se intenta inferir el crecimiento y la viabilidad en el futuro ya que los factores que promueven el establecimiento pueden diferir de aquellos requeridos para la persistencia a largo plazo. Ensamblamos un conjunto grande de datos representativos en general (n = 275) de las reubicaciones de plantas en riesgo bien documentadas y monitoreadas (7.9 años en promedio) para identificar los atributos de sitio más importantes, las técnicas de manejo y los rasgos de las especies para seis parámetros de ciclos de vida y medidas poblacionales del éxito de reubicación. Usamos el algoritmo de bosque aleatorio para cuantificar la importancia relativa de las 29 variables de pronosticadores para cada medida del éxito. Los factores en los resultados de las reubicaciones variaron con los marcos temporales y las medidas de éxito. Las técnicas de manejo tuvieron la mayor influencia relativa sobre la obtención de parámetros de ciclos de vida y tendencias poblacionales a corto plazo, mientras que los atributos de sitio y los rasgos de la especie fueron más importantes para la persistencia poblacional y las tendencias a largo plazo. En específico, las grandes cantidades de fundadores incrementaron el potencial de reproducción y reclutamiento de la siguiente generación, mientras que la declinación de la calidad del hábitat incrementó el riesgo de extinción y el trasplante de especies con baja producción de semillas redujo el rendimiento del potencial reproductivo a la larga. También detectamos interacciones novedosas entre algunos de los factores más importantes, como el aumento en la probabilidad del reclutamiento en la siguiente generación en especies con tasas mayores de producción de semillas, pero sólo cuando se emparejó con grandes cantidades de fundadores. Ya que las barreras más significativas para el éxito de la reubicación de plantas pueden superarse al mejorar las técnicas o resolver los temas a nivel de sitio por medio de un manejo y una intervención temprana, sugerimos que con la combinación del monitoreo a largo plazo con el manejo adaptativo los programas de reubicación pueden aumentar el prospecto de lograr el éxito a largo plazo.

Demographic trade-offs and functional shifts in a hurricane-impacted tropical forest.

BACKGROUND AND AIMS: Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow-fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico. METHODS: We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016. KEY RESULTS: The previously identified growth-survival trade-off was not observed. Instead, we identified a fecundity-growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shift from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time. CONCLUSIONS: Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions.

Costs and benefits of masting: economies of scale are not reduced by negative density-dependence in seedling survival in Sorbus aucuparia.

Masting is a widespread reproductive strategy in plants that helps to reduce seed predation and increase pollination. However, masting can involve costs, notably negative density-dependent (NDD) seedling survival caused by concentrating reproduction in intermittent events. Masting benefits have received widespread attention, but the costs are understudied, which precludes understanding why some plant species have evolved intense masting, while others reproduce regularly. We followed seed production, seed predation (both 13 yr), and seedling recruitment and survival (11 yr) in Sorbus aucuparia. We tested whether NDD in seedling survival after mast years can reduce the benefits of pulsed reproduction that come through predator satiation. Seed predation rates were extreme in our population (mean = 75%), but were reduced by masting. The commonly accepted, but untested, assertion that pulsed recruitment is associated with strong NDD was unsupported. Consequently, the proportion of seedlings that survived their first year increased with fruit production. This provides a rare test of economies of scale beyond the seed stage. Our results provide estimation of the costs of mast seeding, and indicate that these may be lower than expected. Low masting costs, if common, may help explain why masting is such a widespread reproductive strategy throughout the plant kingdom.

Long-term noise pollution affects seedling recruitment and community composition, with negative effects persisting after removal.

Noise pollution can affect species' behaviours and distributions and may hold significant consequences for natural communities. While several studies have researched short-term effects of noise, no long-term research has examined whether observed patterns persist or if community recovery can occur. We used a long-term study system in New Mexico to examine the effects of continuous natural gas well noise exposure on seedling recruitment of foundational tree species (Pinus edulis, Juniperus osteosperma) and vegetation diversity. First, we examined seedling recruitment and vegetation diversity at plots where current noise levels have persisted for greater than 15 years. We then examined recruitment and diversity on plots where noise sources were recently removed or added. We found support for long-term negative effects of noise on tree seedling recruitment, evenness of woody plants and increasingly dissimilar vegetation communities with differences in noise levels. Furthermore, seedling recruitment and plant community composition did not recover following noise removal, possibly due in part to a lag in recovery among animals that disperse and pollinate plants. Our results add to the limited evidence that noise has cascading ecological effects. Moreover, these effects may be long lasting and noise removal may not lead to immediate recovery.

Edge effects alter the role of fungi and insects in mediating functional composition and diversity of seedling recruits in a fragmented tropical forest.

BACKGROUND AND AIMS: In fragmented forests, proximity to forest edges can favour the establishment of resource-acquisitive species over more resource-conservative species. During seedling recruitment, resource-acquisitive species may benefit from either higher light availability or weaker top-down effects of natural enemies. The relative importance of light and enemies for recruitment has seldom been examined with respect to edge effects. METHODS: In a human-modified wet tropical forest in India, we first examined how functional traits indicative of resource-acquisitive vs. resource-conservative strategies, i.e. specific leaf area (SLA), leaf dry matter content, wood density and seed size, explained interspecific differences in densities of seedling recruits with distance to the forest edge. Then, we checked whether fungicide and insecticide treatments and canopy openness (proxy for light availability) explained edge effects on trait-mediated changes in seedling density. Finally, we examined whether light availability and natural enemy activity explained edge effects on functional diversity of seedling recruits. KEY RESULTS: Up to 60 m from edges, recruit densities increased with decreasing seed size, but not at 90-100 m, where recruit densities increased with higher SLA. Trait-mediated variation in recruit densities changed with pesticides only at 90-100 m: compared with control plots, fungicide increased recruit densities for low SLA species and insecticide increased smaller seeded species. For SLA, wood density and seed size, functional diversity of recruits was higher at 90-100 m than at 0-5 m. At 90-100 m, fungicide decreased functional diversity for SLA and insecticide reduced seed size diversity compared with control plots. Canopy openness explained neither variation in recruit density in relation to traits nor functional diversity. CONCLUSIONS: Altered biotic interactions can mediate local changes to trait composition and functional diversity during seedling recruitment in forest fragments, hinting at downstream effects on the structure and function of human-modified forests.

Riparian vegetation model to predict seedling recruitment and restoration alternatives.

Native riparian vegetation communities have declined downstream of large water infrastructure like dams and diversions, owing to water management operations that prevent successful seedling colonization and recruitment. Altered timing and magnitude of reservoir releases to fulfill competing water demands often lead to reduced peak discharges and flow recession rates that do not support native riparian reproduction processes. To achieve short-term ecosystem function in highly regulated rivers an alternative method might be restoration planting, whereby success depends on identifying appropriate planting location and spatial extents. This study aims to provide a methodology to inform resource managers about the extent of possible natural seedling recruitment under average and wet hydrologic conditions, as well as constrain restoration planting operational uncertainties. Results from field surveys and simulations showed limited favorable areas for successful riparian seedling recruitment under regulated flows, regardless of hydrologic conditions in the basin. However, wet (11.4 ha) hydrologic conditions were more (approximately 11 times) favorable than average (1 ha) conditions for seedling recruitment. Furthermore, model results identified the location and spatial extent (25.6 ha) of favorable restoration planting areas during average flow. This extent is approximately 25 times larger than natural recruitment during an average (hydrological) year and even twice that for natural recruitment for a wet year. This suggests that ground operational activities guided by numerical modeling may effectively constrain planting uncertainties.

The timing of frugivore-mediated seed dispersal effectiveness.

The seed dispersal effectiveness framework allows assessing mutualistic services from frugivorous animals in terms of quantity and quality. Quantity accounts for the number of seeds dispersed and quality for the probability of recruitment of dispersed seeds. Research on this topic has largely focused on the spatial patterns of seed deposition because seed fates often vary between microhabitats due to differences in biotic and abiotic factors. However, the temporal dimension has remained completely overlooked despite these factors-and even local disperser assemblages-can change dramatically during long fruiting periods. Here, we test timing effects on seed dispersal effectiveness, using as study case a keystone shrub species dispersed by frugivorous birds and with a fruiting period of 9 months. We evaluated quantity and quality in different microhabitats of a Mediterranean forest and different periods of the fruiting phenophase. We identified the bird species responsible for seed deposition through DNA barcoding and evaluated the probability of seedling recruitment through a series of field experiments on sequential demographic processes. We found that timing matters: The disperser assemblage was temporally structured, seed viability decreased markedly during the plant's fruiting phenophase, and germination was lower for viable seeds dispersed in the fruiting peak. We show how small contributions to seed deposition by transient migratory species can result in a relevant effectiveness if they disperse seeds in a high-quality period for seedling recruitment. This study expands our understanding of seed dispersal effectiveness, highlighting the importance of timing and infrequent interactions for population and community dynamics.

Edge effects reduce α-diversity but not ß-diversity during community assembly in a human-modified tropical forest.

Edge effects can alter the spatial organization of diversity in fragmented habitats. For tropical forests, however, there has been large variation in the strength and direction of such effects reported by different studies. For long-lived organisms like trees, one reason for inconsistent patterns might be due to most studies having examined patterns of diversity and compositional variation in older life stages that bear the legacy of a forest past. Younger life stages can reveal ongoing processes of assembly, but multi-stage examinations are rare. For seedling, sapling, and adult life stages of trees in a human-modified wet tropical forest in the Western Ghats Biodiversity Hotspot (India), we examined how proximity to forest edges (edge influence) modified the spatial organization of diversity. Specifically, for each life stage we tested whether edge influence led to loss of α- and γ-diversity and decreased ß-diversity in this landscape. We found lower α- and γ-diversity closer to forest edges, but only for seedlings. Seedling composition at 90-100 m from forest edges diverged from composition of sites within 60 m, suggesting that edge influence restricted the recruitment of some species to interior sites. In contrast, ß-diversity was greater near edges than interior forest for all life stages and most prominently for seedlings. Furthermore, ß-diversity at edges was primarily driven by species turnover, suggesting either marked species-environment associations or dispersal limitation. Low turnover at 90-100 m implies that ß-diversity arose from stochastic fluctuations in occurrences and abundances of the same species set. Overall, we find that high ß-diversity structured spatial patterns of diversity near edges, but recruitment bottlenecks are likely to reduce alpha diversity of forest fragments. Our results also corroborate the need to maintain sufficiently large areas of tropical forest free from edge effects to avoid the loss of interior forest species. To improve landscape-scale diversity of fragmented landscapes, restoration efforts should focus on recovery of species that are unable to regenerate near forest edges.

Fire cues trigger germination and stimulate seedling growth of grass species from Brazilian savannas.

PREMISE: Although fire cues (high temperatures and smoke) influence seed germination in numerous species from fire-prone environments, their effects on seed germination of species from neotropical savannas are poorly understood. METHODS: We exposed seeds of eight grass species from the Cerrado, the Brazilian savanna to heat-shock (80°C or 110°C for 5 min) and/or smoke water, and then set them to germinate in light or dark, at either summer (28°C/18°C) or winter (27°C/14°C) temperature regimes in an incubator. In addition, we evaluated the effects of smoke water on seedling root and shoot growth for four of the species. RESULTS: Smoke interacted with the dark treatment to increase germination from 28% to 93% in Aristida recurvata and 77% to 95% in Aristida riparia. Smoke had no effect on germination of either of these species in the light. Heat-shock alone also promoted seed germination in A. recurvata. For Digitaria lehmanniana, smoke interacted with heat-shock to improve germination from 5% to 16%. In contrast, the fire treatments did not have any effect on the seed germination of the remaining five species. Smoke water stimulated root growth for A. riparia, A. recurvata, and Ctenium cirrosum but had no effect on their shoot growth. CONCLUSIONS: The strong promotive effect of smoke on Aristida germination suggests that these species are fire-adapted. Aristida species have an active awn system, which facilitates seed burial, and the smoke and dark interaction would ensure buried seeds germinated post-fire. The species that showed no response to fire cues may either have adapted via alternative strategies or require different concentrations of smoke or levels of heat. This study is one of very few examples showing a positive germination and seedling growth response to smoke for species from neotropical savannas.

Effects of life history and reproduction on recruitment time lags in reintroductions of rare plants.

Reintroductions are important components of conservation and recovery programs for rare plant species, but their long-term success rates are poorly understood. Previous reviews of plant reintroductions focused on short-term (e.g., ≤3 years) survival and flowering of founder individuals rather than on benchmarks of intergenerational persistence, such as seedling recruitment. However, short-term metrics may obscure outcomes because the unique demographic properties of reintroductions, including small size and unstable stage structure, could create lags in population growth. We used time-to-event analysis on a database of unusually well-monitored and long-term (4-28 years) reintroductions of 27 rare plant species to test whether life-history traits and population characteristics of reintroductions create time-lagged responses in seedling recruitment (i.e., recruitment time lags [RTLs]), an important benchmark of success and indicator of persistence in reintroduced populations. Recruitment time lags were highly variable among reintroductions, ranging from <1 to 17 years after installation. Recruitment patterns matched predictions from life-history theory with short-lived species (fast species) exhibiting consistently shorter and less variable RTLs than long-lived species (slow species). Long RTLs occurred in long-lived herbs, especially in grasslands, whereas short RTLs occurred in short-lived subtropical woody plants and annual herbs. Across plant life histories, as reproductive adult abundance increased, RTLs decreased. Highly variable RTLs were observed in species with multiple reintroduction events, suggesting local processes are just as important as life-history strategy in determining reintroduction outcomes. Time lags in restoration outcomes highlight the need to scale success benchmarks in reintroduction monitoring programs with plant life-history strategies and the unique demographic properties of restored populations. Drawing conclusions on the long-term success of plant reintroduction programs is premature given that demographic processes in species with slow life-histories take decades to unfold.

Efectos de la Historia de Vida y la Reproducción sobre las Demoras en el Tiempo de Reclutamiento en la Reintroducción de Plantas Raras Resumen Las reintroducciones son componentes importantes de los programas de conservación y recuperación de especies raras de plantas, pero las tasas de éxito a largo plazo cuentan con muy poco entendimiento. Las revisiones previas de las reintroducciones de plantas se han enfocado en la supervivencia a corto plazo (p. ej.: ≤ 3 años) y en el florecimiento de individuos fundadores en lugar de enfocarse en puntos de referencia para la persistencia inter-generacional, como el reclutamiento de plántulas. Sin embargo, las medidas a corto plazo pueden ocultar los resultados ya que las propiedades demográficas únicas de las reintroducciones, incluyendo el menor tamaño y la estructura inestable de estadio, podrían crear demoras en el crecimiento poblacional. Usamos un análisis de tiempo-para-evento en una base de datos de reintroducciones inusualmente bien monitoreadas y de largo plazo (4-28 años) de 27 especies raras de plantas para probar si los atributos de la historia de vida y las características poblacionales de la reintroducción crean respuestas con demoras temporales en el reclutamiento de plántulas (es decir, demoras temporales en el reclutamiento), un punto de referencia importante para el éxito y un indicador de la persistencia en poblaciones reintroducidas. Las demoras temporales de reclutamiento (RTLs, en inglés) fueron muy variables entre las reintroducciones, abarcando desde <1 hasta 17 años después de la instalación. Los patrones de reclutamiento se acoplaron a las predicciones de la teoría de historias de vida, donde las especies de vida corta (especies rápidas) exhibieron RTLs consistentemente más cortas y menos variables que las especies de vida larga (especies lentas). Las RTLs largas ocurrieron en hierbas de vida larga, especialmente en los pastizales, mientras que las RTLs cortas ocurrieron en plantas leñosas subtropicales de vida corta y en hierbas anuales. En todas las historias de vida de las plantas, conforme incrementó la abundancia de adultos reproductivos, las RTLs disminuyeron. Se observaron RTLs altamente variables en las especies con eventos de reintroducción múltiples, lo que sugiere que los procesos locales son igual de importantes que la estrategia de historia de vida para determinar los resultados de las reintroducciones. Las demoras temporales en los resultados de restauración resaltan la necesidad de poner a escala los puntos de referencia de éxito en los programas de monitoreo de reintroducciones que tengan estrategias de historia de vida de las plantas y las propiedades demográficas únicas de las poblaciones restauradas. La obtención de conclusiones sobre el éxito a largo plazo de los programas de reintroducción de plantas es algo prematuro ya que los procesos demográficos de especies con historias de vida lentas tardan décadas en desarrollarse.

Inorganic soil amendments alter seedling performance of native plant species in post-mining arid zone rehabilitation.

Rehabilitation of degraded drylands is challenged by environmental and anthropogenic constraints, such as limited availability of locally-sourced topsoil and poor quality alternative soil substrates. Current rehabilitation practices, at times, utilise inorganic soil amendments to improve the physicochemical and biological characteristics of reconstructed soil profiles. These approaches may be appropriate for dryland rehabilitation, but there is limited research available regarding the benefits of using these amendments. Here, we present a study in the Pilbara region of Western Australia, an arid landscape subject to intensive mining that currently uses inorganic soil amendments (gypsum and urea) in post-mining rehabilitation. The aim of this study was to assess the effectiveness of these amendments to (1) promote seed germination, seedling emergence and seedling growth across five plant species and, (2) re-instate soil quality in mine waste substrates. A series of glasshouse experiments assessed eight application combinations of these amendments in two alternative substrates and compared these to unamended substrates and topsoil. Soil amendments had a limited influence on seed germination, were detrimental to seedling emergence and resulted in increased seedling mortality. Mortality in the waste ranged from 2 to 61% but increased to 7-92% in amended waste. Seedling growth improved with high doses of amendments in waste, with a 1.3-5.6-fold increase across all plant species. Soil quality was relatively unaffected by amendments with soil nitrogen ranging from 0.01 to 0.08%, organic carbon from 0.01 to 0.12% and soil microbial activity from 2.3 to 2.4 ppm-CO2 in the amended and unamended waste. The use of soil amendments in mine rehabilitation requires consideration of the trade-off between initial reductions in seedling recruitment and enhanced seedling development at later stages. Future rehabilitation should consider the timing of amendment application to avoid detrimental impacts on seedling recruitment and maximise the benefits to seedling growth.

High land-use intensity exacerbates shifts in grassland vegetation composition after severe experimental drought.

Climate change projections anticipate increased frequency and intensity of drought stress, but grassland responses to severe droughts and their potential to recover are poorly understood. In many grasslands, high land-use intensity has enhanced productivity and promoted resource-acquisitive species at the expense of resource-conservative ones. Such changes in plant functional composition could affect the resistance to drought and the recovery after drought of grassland ecosystems with consequences for feed productivity resilience and environmental stewardship. In a 12-site precipitation exclusion experiment in upland grassland ecosystems across Switzerland, we imposed severe edaphic drought in plots under rainout shelters and compared them with plots under ambient conditions. We used soil water potentials to scale drought stress across sites. Impacts of precipitation exclusion and drought legacy effects were examined along a gradient of land-use intensity to determine how grasslands resisted to, and recovered after drought. In the year of precipitation exclusion, aboveground net primary productivity (ANPP) in plots under rainout shelters was -15% to -56% lower than in control plots. Drought effects on ANPP increased with drought severity, specified as duration of topsoil water potential ψ < -100 kPa, irrespective of land-use intensity. In the year after drought, ANPP had completely recovered, but total species diversity had declined by -10%. Perennial species showed elevated mortality, but species richness of annuals showed a small increase due to enhanced recruitment. In general, the more resource-acquisitive grasses increased at the expense of the deeper-rooted forbs after drought, suggesting that community reorganization was driven by competition rather than plant mortality. The negative effects of precipitation exclusion on forbs increased with land-use intensity. Our study suggests a synergistic impact of land-use intensification and climate change on grassland vegetation composition, and implies that biomass recovery after drought may occur at the expense of biodiversity maintenance.

Adaptive differentiation among populations of the Mediterranean dry grassland species Brachypodium retusum: The role of soil conditions, grazing, and humidity.

PREMISE OF THE STUDY: Genetic differentiation in plant species may result from adaptation to environmental conditions, but also from stochastic processes. The drivers selecting for local adaptation and the contribution of adaptation to genetic differentiation are often unknown. Restoration and succession studies have revealed different colonization patterns for Brachypodium retusum, a common Mediterranean grass. In order to understand these patterns, we tested population differentiation and adaptation to different environmental factors. METHODS: Structured sampling of 12 populations from six sites and two soil types within site was used to analyze the spatial and environmental structure of population differentiation. Sampling sites differ in grazing intensity and climate. We tested germination and growth in a common garden. In subsets, we analyzed the differential response to stone cover, grazing and soil moisture. KEY RESULTS: We found significant differences among populations. The site explained population differentiation better than soil, suggesting a dominant influence of climate and/or genetic drift. Stone cover had a positive influence on seedling establishment, and populations showed a differential response. However, this response was not related to environmental differences between collection sites. Regrowth after clipping was higher in populations from the more intensively grazed Red Mediterranean soils suggesting an adaptation to grazing. Final germination was generally high even under drought, but germination response to differences in soil moisture was similar across populations. CONCLUSIONS: Adaptive population differentiation in germination and early growth may have contributed to different colonization patterns. Thus, the provenance of B. retusum needs to be carefully considered in ecological restoration.

Selective predation on acorn weevils by seed-caching Siberian chipmunk Tamias sibiricus in a tripartite interaction.

Although food-hoarding animals benefit plant seeds by generating predation pressure on granivorous insects, we lack experimental evidence of whether the tripartite interactions maintain a mutualistic relationship between the third trophic level and primary producers. Relying on the behavior of shelling, Siberian chipmunks (Tamias sibiricus) selectively consumed weevil larvae infested in acorns of Mongolian oak (Quercus mongolica) but chose the non-infested acorns to scatter-hoard. Shelling not only reduced volatile emission from acorns but also decreased cache loss to pilferers, weevil larvae and fungi, allowing T. sibiricus to gain more rewards from their caches. Moreover, shelling by T. sibiricus enhanced acorn germination and seedling establishment of Q. mongolica, possibly due to the diminishment of the negative effects of weevil infestation on acorn viability. Here, we show that both food-hoarding animal T. sibiricus and oak Q. mongolica can be conditionally benefited from selective predation on weevil larvae inside acorns. Our results highlight the need to integrate the mutualisms between the third and first trophic level into the tripartite interaction model. We suggest that more efforts should be made in the tripartite interactions of food-hoarding animals, seeds and granivorous insects.

Frequency-dependent selection for rare genotypes promotes genetic diversity of a tropical palm.

Negative frequency-dependent selection among species is a key driver of community diversity in natural systems, but the degree to which negative frequency-dependent selection shapes patterns of survival and genetic diversity within species is poorly understood. In a 5-year field experiment, we show that seedlings of a tropical palm with rare genotypes had a pronounced survival advantage over seedlings with common genotypes, with effect sizes comparable to that of light availability. This 'rare genotype advantage' led to an increase in population-wide genetic diversity among seedlings compared to null expectations, as predicted by negative frequency-dependent selection, and increased reproductive success in adult trees with rare genotypes. These results suggest that within-species negative frequency-dependent selection of genotypes can shape genetic variation on ecologically relevant timescales in natural systems and may be a key, overlooked source of non-random mortality for tropical plants.

Seed-bank structure and plant-recruitment conditions regulate the dynamics of a grassland-shrubland Chihuahuan ecotone.

Large areas of desert grasslands in the southwestern United States have shifted to sparse shrublands dominated by drought-tolerant woody species over the last 150 yr, accompanied by accelerated soil erosion. An important step toward the understanding of patterns in species dominance and vegetation change at desert grassland-shrubland transitions is the study of environmental limitations imposed by the shrub-encroachment phenomenon on plant establishment. Here, we analyze the structure of soil seed banks, environmental limitations for seed germination (i.e., soil-water availability and temperature), and simulated seedling emergence and early establishment of dominant species (black grama, Bouteloua eriopoda, and creosotebush, Larrea tridentata) across a Chihuahuan grassland-shrubland ecotone (Sevilleta National Wildlife Refuge, New Mexico, USA). Average viable seed density in soils across the ecotone is generally low (200-400 seeds/m2 ), although is largely concentrated in densely vegetated areas (with peaks up to 800-1,200 seeds/m2 in vegetated patches). Species composition in the seed bank is strongly affected by shrub encroachment, with seed densities of grass species sharply decreasing in shrub-dominated sites. Environmental conditions for seed germination and seedling emergence are synchronized with the summer monsoon. Soil-moisture conditions for seedling establishment of B. eriopoda take place with a recurrence interval ranging between 5 and 8 yr for grassland and shrubland sites, respectively, and are favored by strong monsoonal precipitation. Limited L. tridentata seed dispersal and a narrow range of rainfall conditions for early seedling establishment (50-100 mm for five to six consecutive weeks) constrain shrub-recruitment pulses to localized and episodic decadal events (9-25 yr recurrence intervals) generally associated with late-summer rainfall. Re-establishment of B. eriopoda in areas now dominated by L. tridentata is strongly limited by the lack of seeds and decreased plant-available soil moisture for seedling establishment.

Interspecific associations in seed arrival and seedling recruitment in a Neotropical forest.

Contagious seed dispersal refers to the tendency for some sites to receive many dispersed seeds while other sites receive few dispersed seeds. Contagious dispersal can lead to interspecific associations in seed arrival, and this in turn might lead to interspecific associations in seedling recruitment. We evaluate the extent of spatially contagious seed arrival, the frequency of positive interspecific associations in seed arrival, and their consequences for seedling recruitment at the community level in a tropical moist forest. We quantified seed arrival to 200 passive seed traps for 28 yr of weekly censuses and seedling recruitment to 600 1-m2 quadrats for 21 yr of annual censuses on Barro Colorado Island, Panama. We assessed whether spatially contagious seed dispersal was more important among zoochorous species than among anemochorous species, increased in importance with similarity in fruiting times, and led to interspecific associations in seed arrival and seedling recruitment. We controlled adult seed source associations statistically to evaluate predicted relationships. We found that spatially contagious seed arrival was widespread among zoochorous species, but also occurred among anemochorous species when the strong, consistent trade winds were present. Significant interspecific associations in seed arrival were more likely for pairs of species with zoochorous seeds and similar fruiting times and persisted through seedling recruitment. Thus, interspecifically contagious seed dispersal affects local species composition and alters the mixture of interspecific interactions through the seed, germination, and early seedling stages in this forest. Future investigations should consider the implications of interspecific association at the regeneration stages documented here for later life stages and species coexistence.

Climate warming could increase recruitment success in glacier foreland plants.

BACKGROUND AND AIMS: Glacier foreland plants are highly threatened by global warming. Regeneration from seeds on deglaciated terrain will be crucial for successful migration and survival of these species, and hence a better understanding of the impacts of climate change on seedling recruitment is urgently needed to predict future plant persistence in these environments. This study presents the first field evidence of the impact of climate change on recruitment success of glacier foreland plants. METHODS: Seeds of eight foreland species were sown on a foreland site at 2500 m a.s.l., and at a site 400 m lower in altitude to simulate a 2·7 °C increase in mean annual temperature. Soil from the site of origin was used to reproduce the natural germination substrate. Recruitment success, temperature and water potential were monitored for 2 years. The response of seed germination to warming was further investigated in the laboratory. KEY RESULTS: At the glacier foreland site, seedling emergence was low (0 to approx. 40 %) and occurred in summer in all species after seeds had experienced autumn and winter seasons. However, at the warmer site there was a shift from summer to autumn emergence in two species and a significant increase of summer emergence (13-35 % higher) in all species except two. Survival and establishment was possible for 60-75 % of autumn-emerged seedlings and was generally greater under warmer conditions. Early snowmelt in spring caused the main ecological factors enhancing the recruitment success. CONCLUSIONS: The results suggest that warming will influence the recruitment of glacier foreland species primarily via the extension of the snow-free period in spring, which increases seedling establishment and results in a greater resistance to summer drought and winter extremes. The changes in recruitment success observed here imply that range shifts or changes in abundance are possible in a future warmer climate, but overall success may be dependent on interactions with shifts in other components of the plant community.

Germination and seedling establishment in orchids: a complex of requirements.

BACKGROUND: Seedling recruitment is essential to the sustainability of any plant population. Due to the minute nature of seeds and early-stage seedlings, orchid germination in situ was for a long time practically impossible to observe, creating an obstacle towards understanding seedling site requirements and fluctuations in orchid populations. The introduction of seed packet techniques for sowing and retrieval in natural sites has brought with it important insights, but many aspects of orchid seed and germination biology remain largely unexplored. KEY CONSIDERATIONS: The germination niche for orchids is extremely complex, because it is defined by requirements not only for seed lodging and germination, but also for presence of a fungal host and its substrate. A mycobiont that the seedling can parasitize is considered an essential element, and a great diversity of Basidiomycota and Ascomycota have now been identified for their role in orchid seed germination, with fungi identifiable as imperfect Rhizoctonia species predominating. Specificity patterns vary from orchid species employing a single fungal lineage to species associating individually with a limited selection of distantly related fungi. A suitable organic carbon source for the mycobiont constitutes another key requirement. Orchid germination also relies on factors that generally influence the success of plant seeds, both abiotic, such as light/shade, moisture, substrate chemistry and texture, and biotic, such as competitors and antagonists. Complexity is furthermore increased when these factors influence seeds/seedling, fungi and fungal substrate differentially. CONCLUSIONS: A better understanding of germination and seedling establishment is needed for conservation of orchid populations. Due to the obligate association with a mycobiont, the germination niches in orchid species are extremely complex and varied. Microsites suitable for germination can be small and transient, and direct observation is difficult. An experimental approach using several levels of environmental manipulation/control is recommended.