Forests are not immune to plant invasions: phenotypic plasticity and local adaptation allow <Emphasis Type="Italic">Prunella vulgaris</Emphasis> to colonize a temperate evergreen rainforest

In the South American temperate evergreen rainforest (Valdivian forest), invasive plants are mainly restricted to open sites, being rare in the shaded understory. This is consistent with the notion of closed-canopy forests as communities relatively resistant to plant invasions. However, alien plants able to develop shade tolerance could be a threat to this unique forest. Phenotypic plasticity and local adaptation are two mechanisms enhancing invasiveness. Phenotypic plasticity can promote local adaptation by facilitating the establishment and persistence of invasive species in novel environments. We investigated the role of these processes in the recent colonization of Valdivian forest understory by the perennial alien herb Prunella vulgaris from nearby populations in open sites. Using reciprocal transplants, we found local adaptation between populations. Field data showed that the shade environment selected for taller plants and greater specific leaf areas. We found population differentiation and within-population genetic variation in both mean values and reaction norms to light variation of several ecophysiological traits in common gardens from seeds collected in sun and shade populations. The colonization of the forest resulted in a reduction of plastic responses to light variation, which is consistent with the occurrence of genetic assimilation and suggests that P. vulgaris individuals adapted to the shade have reduced probabilities to return to open sites. All results taken together confirm the potential for rapid evolution of shade tolerance in P. vulgaris and suggest that this alien species may pose a threat to the native understory flora of Valdivian forest.     

Soil seed banks in different environments: initial forest,mature forest,Pinus and Eucalyptus abandoned stands

Abstract

Different environments (initial forest, mature forest, Pinus and Eucalyptus stands) found in Seasonal Semideciduous Forest fragments affect the density of viable seeds, as well as the floristic similarity, diversity, and richness of tree species in soil seed banks. This hypothesis was tested in the current study. Soil seed bank samples were collected in the aforementioned environments during rainy and dry seasons, and taken to a shade house, where they remained under favorable seed germination conditions. Tree seedling emergence was measured, and sample species were identified every 15?days, for six months, in each sampling period. In total, 97 individuals m^?2 and 23 species emerged in all environments and periods. The highest density of viable seeds of tree species in the soil seed bank was found in the initial forest stretch, mature forest stretch and abandoned Eucalyptus stand. Only the Pinus stand seed bank in the dry season had different floristic and lower viable seed density than the mature forest seed bank. Thus, all environments, except the abandoned Pinus stand, can preserved Seasonal Semideciduous Forest fragments.     

Which Extent is Plasticity to Light Involved in the Ecotypic Differentiation of a Tree Species from Savanna and Forest?

Light intensity and heterogeneity are some of the main environmental factors that differ between forest and savanna habitats, and plant species from these habitats form distinct functional types. In this study, we tested the hypothesis that not only differences in morphological and physiological traits but also phenotypic plasticity in response to light are involved in adaptation to forest and savanna habitats by investigating ecotypic differentiation between populations of Plathymenia reticulata (Leguminosae: Mimosoideae), a tree from the Brazilian Atlantic Forest and the Brazilian Cerrado (savanna). Seeds from four natural populations (one from each biome core area and two from ecotonal regions) were grown in a common garden with four light treatments. Fifteen morphological and physiological characteristics were evaluated until individuals reached 6 mo old. Comparisons among populations showed differences for seven traits in at least one light treatment. These differences pointed to local adaptation to different biomes. Populations showed different levels of phenotypic plasticity in response to light in seven traits. Higher plasticity was found either in the forest core population or ecotonal populations; lower values were found in the cerrado core population. Lower plasticity in the cerrado population emphasizes the stress resistant syndrome, as lower plasticity is probably advantageous in a habitat where a conservative resource use is crucial. Higher plasticity in forest individuals suggests higher ability in exploiting the light heterogeneity in this habitat. Also, higher plasticity in ecotonal populations can be important to ensure the maintenance of P. reticulata in these temporally and spatially dynamic areas. Abstract in Portugese is available at http://www.blackwell‐synergy.com/loi/btp .     

Differences in developmental strategies between long‐settled and invasion‐front populations of the cane toad in Australia

Phenotypic plasticity can enhance a species’ ability to persist in a new and stressful environment, so that reaction norms are expected to evolve as organisms encounter novel environments. Biological invasions provide a robust system to investigate such changes. We measured the rates of early growth and development in tadpoles of invasive cane toads (Rhinella marina) in Australia, from a range of locations and at different larval densities. Populations in long‐colonized areas have had the opportunity to adapt to local conditions, whereas at the expanding range edge, the invader is likely to encounter challenges that are both novel and unpredictable. We thus expected invasion‐vanguard populations to exhibit less phenotypic plasticity than range‐core populations. Compared to clutches from long‐colonized areas, clutches from the invasion front were indeed less plastic (i.e. rates of larval growth and development were less sensitive to density). In contrast, those rates were highly variable in clutches from the invasion front, even among siblings from the same clutch under standard conditions. Clutches with highly variable rates of growth and development under constant conditions had lower phenotypic plasticity, suggesting a trade‐off between these two strategies. Although these results reveal a strong pattern, further investigation is needed to determine whether these different developmental strategies are adaptive (i.e. adaptive phenotypic plasticity vs. bet‐hedging) or instead are driven by geographic variation in genetic quality or parental effects.     

Patterns of Differentiation in Wiregrass (Aristida beyrichiana): Implications for Restoration Efforts

Aristida beyrichiana (wiregrass) is increasingly being planted in restoration projects across the southeastern coastal plain, with little focus on genetic differences among populations across the region. Local and regional population differentiation for establishment and growth traits were examined in common garden and reciprocal transplant experiments. Seeds from up to 20 plants from each of seven populations were collected in northern and central Florida sites that encompassed gradients of soils, hydrology, and temperature. Reciprocal seed transplants using three of the common garden populations were conducted in two consecutive years. In the common garden, significant population differences were seen in seed weight, seedling emergence and survival, tiller height, number of tillers, the relationship between tiller number and tiller height, and flowering. Variation among maternal families was seen in tiller number and in the relationship between tiller number and tiller height. The reciprocal transplant study did not detect either local adaptation to sites of origin or consistent superiority of one source population or planting site in seedling establishment. These results suggest that the probability of seedling establishment is primarily dependent on environmental conditions rather than genetic differences. Genetic variation for traits related to fitness (e.g., tiller number) may be retained within populations because phenotypically plastic growth responses of seedlings to environmental variation buffer genetic variation against the action of selection. But despite the lack of evidence for genetic influences on initial establishment in wiregrass, our common garden study suggests genetic differences among populations. This result, when combined with previous results indicating local adaptation in later life stages of wiregrass, suggests that restoration efforts involving this species should use local seed sources from sites with similar soil and hydrological conditions.     

Large herbivores regulate the spatial recruitment of a hyperdominant Neotropical palm

Large mammalian herbivores play an important role in shaping the diversity of tropical forests by affecting the survival of seedlings and saplings beneath parent plants. The white‐lipped peccary (Tayassu pecari) accounts for the largest herbivore biomass that controls seed and seedling survival in Neotropical ecosystems. However, hunting and habitat loss has driven peccaries to local extinction for most of their original distribution, so it is likely that their absence will affect plant recruitment dynamics. We tested the effects of peccary local extinction on the density and spatial distribution of the hyperdominant palm Euterpe edulis by performing a fine‐scale characterization of its spatial recruitment in six forest sites in the Brazilian Atlantic forest. We compared the age structure and the spatial patterns of seedlings, saplings, and adults as well as the relationship between them. We found that while under the presence of peccaries there was a decrease in recruitment rates under adults, the local extinction of these large mammals led to a more clumped process of spatial recruitment. Despite such contrasting spatial patterns of recruitment dynamics, neither age structure nor the random spatial distribution of adults was affected by the presence or absence of peccaries, indicating that their early effects on these palm populations are mitigated as recruitment advances. Our findings highlight the role of large‐bodied forest‐dwelling herbivores in regulating the fine‐scale spatial recruitment of plants and advance our understanding on the effects of defaunation in tropical forests. Abstract in Portuguese is available with online material.     

Local Adaptation and Effects of Grazing among Seedlings of Two Native California Bunchgrass Species: Implications for Restoration

Adaptation to environmental factors may influence the germination and establishment of focal species in ecological restoration. Reciprocal transplants remain one of the best methods to detect local adaptation, but long‐term studies are often not feasible. We conducted reciprocal transplants of the native California bunchgrasses Elymus glaucus and Bromus carinatus between two central California locations to seek evidence of adaptation to local environmental conditions in a single growing season. Experimental plots at one location included grazed and ungrazed sites. The combination of locations and grazing treatments allowed us to determine whether the ability to detect evidence for adaptation depended on grazing regime. In addition, we measured the direct effects of grazing on seedling growth and survival concurrent with our investigation of local adaptation. We detected a homesite advantage for seedling growth or survival in both species, but the factors contributing to adaptive differentiation were species specific. Evidence of local adaptation was detected for seedling biomass in Bromus and for survivorship in Elymus. The homesite advantage observed in both species was greatly reduced under grazed conditions and in Elymus was significant only in the ungrazed plots. Climate and soil analyses detected significant differences between locations in five soil attributes and two climate variables. In particular, differences in exchangeable magnesium indicated that one of the two transplant locations consisted of serpentine soil, which is widely known to drive adaptation in plant populations. Together, these results suggest that it is possible to investigate the scale and factors involved in local adaptation with short‐term transplant studies.     

Is <Emphasis Type="Italic">Pinus radiata</Emphasis> invading the native vegetation in central Chile? Demographic responses in a fragmented forest

Forest fragmentation facilitates the invasion of exotic species. This threat may be especially severe if forest fragments are surrounded by plantations of exotic species like Pinus radiata, an aggressive colonizer and shade-intolerant tree that has invaded successfully several native ecosystems of the southern hemisphere. In this study, we experimentally tested if the conditions of a successful seedling establishment P. radiata are fulfilled at the Coastal Maulino forest, an endemic fragmented forest of central Chile. Results demonstrated that seeds are dispersed into the native forests, however seedling establishment occurs only at the edges. We conclude that this exotic species is not invading native forests up to date. However, we suggest to conduct evaluations of seed rain and seedling establishment in the long term, in order to monitor the fate of this exotic species in fragmented native forest of Central Chile.     

Landscape‐scale deforestation decreases gene flow distance of a keystone tropical palm,Euterpe edulis Mart (Arecaceae)

Habitat loss represents one of the main threats to tropical forests, which have reached extremely high rates of species extinction. Forest loss negatively impacts biodiversity, affecting ecological (e.g., seed dispersal) and genetic (e.g., genetic diversity and structure) processes. Therefore, understanding how deforestation influences genetic resources is strategic for conservation. Our aim was to empirically evaluate the effects of landscape‐scale forest reduction on the spatial genetic structure and gene flow of Euterpe edulis Mart (Arecaceae), a palm tree considered a keystone resource for many vertebrate species. This study was carried out in nine forest remnants in the Atlantic Forest, northeastern Brazil, located in landscapes within a gradient of forest cover (19–83%). We collected leaves of 246 adults and 271 seedlings and performed genotyping using microsatellite markers. Our results showed that the palm populations had low spatial genetic structure, indicating that forest reduction did not influence this genetic parameter for neither seedlings nor adults. However, forest loss decreased the gene flow distance, which may negatively affect the genetic diversity of future generations by increasing the risk of local extinction of this keystone palm. For efficient strategies of genetic variability conservation and maintenance of gene flow in E. edulis, we recommend the maintenance of landscapes with intermediary to high levels of forest cover, that is, forest cover above 40%.     

Exploiting fruits of a threatened palm to trigger restoration of Brazil's Atlantic Forest

Recent global commitments to forest and landscape restoration in the tropics call for new management approaches that benefit both biodiversity and livelihoods of forest‐dependent people. The sustainable use of wild forest products is a promising pathway, but requires clarity about harvested species' demography and harvesters' rights. Here, we explored how the exploitation of fruits of the threatened palm Euterpe edulis, a key fruit source for wildlife in Brazil's globally important Atlantic Forest biodiversity hotspot, could trigger local community involvement in restoration. This palm has both non‐destructive (fruit) and destructive (palm heart) culinary uses, each with unique biological and resource‐use conditions. We quantified all demographic stages of this species in multiple agroforests, secondary forests, and protected areas to parameterize demographic projections of palm populations, harvest profitability, and fruit provisioning to wildlife under different management scenarios. Field observations showed a clear depression of adult palm populations in protected areas, likely due to palm heart poaching, and of intermediate size classes in agroforests, probably from weeding by farmers. Field data and demographic models reveal that in this region, agroforests and secondary forests can successfully conserve this species while providing lasting profits for farmers under most scenarios. These findings demonstrate a clear case where local stakeholders both contribute to and benefit from restoration through harvest of wild products and also highlight a potential source of income from regenerating tropical forests.     

Phenotypic plasticity closely linked to climate at origin and resulting in increased mortality under warming and frost stress in a common grass

Phenotypic plasticity is important for species responses to global change and species coexistence. Phenotypic plasticity differs among species and traits and changes across environments. Here, we investigated phenotypic plasticity of the widespread grass Arrhenatherum elatius in response to winter warming and frost stress by comparing phenotypic plasticity of 11 geographically and environmentally distinct populations of this species to phenotypic plasticity of populations of different species originating from a single environment. The variation in phenotypic plasticity was similar for populations of a single species from different locations compared to populations of functionally and taxonomically diverse species from one environment for the studied traits (leaf biomass production and root integrity after frost) across three indices of phenotypic plasticity (RDPI, PIN, slope of reaction norm). Phenotypic plasticity was not associated with neutral genetic diversity but closely linked to the climate of the populations’ origin. Populations originating from warmer and more variable climates showed higher phenotypic plasticity. This indicates that phenotypic plasticity can itself be considered as a trait subject to local adaptation to climate. Finally, our data emphasize that high phenotypic plasticity is not per se positive for adaptation to climate change, as differences in stress responses are resulting in high phenotypic plasticity as expressed by common plasticity indices, which is likely to be related to increased mortality under stress in more plastic populations.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

A mid‐Pleistocene rainforest corridor enabled synchronous invasions of the Atlantic Forest by Amazonian anole lizards

Shifts in the geographic distribution of habitats over time can promote dispersal and vicariance, thereby influencing large‐scale biogeographic patterns and ecological processes. An example is that of transient corridors of suitable habitat across disjunct but ecologically similar regions, which have been associated with climate change over time. Such connections likely played a role in the assembly of tropical communities, especially within the highly diverse Amazonian and Atlantic rainforests of South America. Although these forests are presently separated by open and dry ecosystems, paleoclimatic and phylogenetic evidence suggest that they have been transiently connected in the past. However, little is known about the timing, magnitude and the distribution of former forest connections. We employ sequence data at multiple loci from three codistributed arboreal lizards (Anolis punctatus, Anolis ortonii and Polychrus marmoratus) to infer the phylogenetic relationships among Amazonian and Atlantic Forest populations and to test alternative historical demographic scenarios of colonization and vicariance using coalescent simulations and approximate Bayesian computation (ABC). Data from the better‐sampled Anolis species support colonization of the Atlantic Forest from eastern Amazonia. Hierarchical ABC indicates that the three species colonized the Atlantic Forest synchronously during the mid‐Pleistocene. We find support of population bottlenecks associated with founder events in the two Anolis, but not in P. marmoratus, consistently with their distinct ecological tolerances. Our findings support that climatic fluctuations provided key opportunities for dispersal and forest colonization in eastern South America through the cessation of environmental barriers. Evidence of species‐specific histories strengthens assertions that biological attributes play a role in responses to shared environmental change.     

Assessing the effect of genetic diversity on the early establishment of the threatened seagrass Posidonia australis using a reciprocal‐transplant experiment

Two common goals for restoration are rapid plant establishment and long‐term plant persistence. The success of transplanted populations may be jeopardized if the donor transplants are not genetically diverse, and/or poorly matched to their new environment. Here, we test the effects of local adaptation and plot‐level genetic diversity on the early establishment phase of a threatened seagrass species, Posidonia australis, by performing a reciprocal transplant experiment across two genetically and geographically distinct populations in southeastern Australia. Posidonia australis is a long‐lived, slow‐growing species that has no seed bank, and the successful transplantation of live shoots and seedlings is the only available restoration method. Our results show a strong effect of local adaptation and genetic diversity on P. australis survivorship and performance over the first 6 months following transplantation. High‐genetic diversity plots displayed higher survival rates and exhibited reduced productivity and increased carbohydrate reserves within the rhizome. This suggests that high‐diversity plots included shoots that were conserving energy stores by actively reducing growth rates during the early stages of transplantation. The lowest diversity plots exhibited high leaf and root productivity and corresponding low carbohydrate reserves. This may be a sign of stress in the low‐diversity transplants, potentially explaining the very low survival rate. We suggest that future restoration efforts source donor transplants from multiple local sources to ensure both local adaptation and sufficient genetic diversity to increase the likelihood of early establishment success.     

The response of forest plant regeneration to temperature variation along a latitudinal gradient

Background and Aims

The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming.

Methods

Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments.

Key Results

Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted.

Conclusions

Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.     

Interactions between mountain birch seedlings from differentiated populations in contrasting environments of subarctic Russia

So far very few experiments have accounted for the combined effect of two phenomena co-occurring in stress gradients: local adaptation to stress and the increase in facilitation with increasing stress (predicted by the stress-gradient hypothesis, SGH). Mountain birch (Betula pubescens subsp. czerepanovii) facilitates conspecific seedlings in subarctic high stress sites and is capable of rapid evolutionary adaptation, being therefore a good model species for a study combining local ecotypes and SGH. A within-species experiment was conducted to test SGH in three stress gradients, detect potential local adaptations between low and high stress populations, and assess their effects on seedling-seedling interactions. Although no evidence for local adaptation was detected, high and low stress populations showed some differentiation, possibly explained by decreasing phenotypic plasticity in high stress conditions and/or neutral evolutionary mechanisms. Weak support for SGH was detected. While facilitation was unaffected by seedling origin, low stress populations showed better competitive ability.     

Distribution of Leguminosae tree species in different altitudinal levels along the Atlantic Rain Forest in the Brazilian coast

The Atlantic Forest is one of the most threatened tropical forests in the world. Leguminosae, by its great richness and dominance among arboreal stratum elements, is of major importance in the floristic composition and structure of this forest. We investigated the distribution of legume species on an altitudinal gradient to find out the altitudinal zones with higher richness of species; the altitudinal zones with greater floristic similarity; the possible presence of species that may be exclusive to certain vegetation types and the altitudinal amplitudes of those species, as well as the occurrence of species substitution along the altitudinal gradient. Therefore, thirty one studies conducted in different altitudinal levels between 5° S and 29° S were analyzed. A matrix with 142 tree species distributed in altitudinal zones (every 100 m) from sea level to 2100 m was built. The greatest species richness was observed in the Submontane Forest (50–500 m) with 92 species. The cluster analysis revealed a strong dissimilarity of the 1400–2100 m (Upper Montane) and 0–10 m zones (Restinga Forest). The Submontane and the Montane Forest share the highest number of species (38 ssp.). Forty species are unique to Submontane. Substitution of species was verified. Some species have their preferred habitat located at a specific altitudinal amplitude, as is the case of Inga laurina and I. subnuda (0–10 m), I. lanceifolia and Machaerium scleroxylon (800–1200 m). The Leguminosae, although well adapted to the first colonization and establishment of diverse environment, was poorly represented above 1500 m altitude.     

Mahonia invasions in different habitats: local adaptation or general-purpose genotypes?

Rapid evolutionary adaptations and phenotypic plasticity have been suggested to be two important, but not mutually exclusive, mechanisms contributing to the spread of invasive species. Adaptive evolution in invasive plants has been shown to occur at large spatial scales to different climatic regions, but local adaptation at a smaller scale, e.g. to different habitats within a region, has rarely been studied. Therefore, we performed a case study on invasive Mahonia populations to investigate whether local adaptation may have contributed to their spread. We hypothesized that the invasion success of these populations is promoted by adaptive differentiation in response to local environmental conditions, in particular to the different soils in these habitats. To test this hypothesis, we carried out a reciprocal transplantation experiment in the field using seedlings from five Mahonia populations in Germany that are representative for the range of habitats invaded, and a greenhouse experiment that specifically compared the responses to the different soils of these habitats. We found no evidence for local adaptation of invasive Mahonia populations because seedlings from all populations responded similarly to different habitats and soils. In a second greenhouse experiment we examined genetic variation within populations, but seedlings from different maternal families did not vary in their responses to soil conditions. We therefore suggest that local adaptation of seedlings does not play a major role for the invasion success of Mahonia populations and that phenotypic plasticity, instead, could be an important trait in this stage of the life cycle.     

Phenotypic plasticity in sex allocation for a simultaneously hermaphroditic coral reef fish

Phenotypic plasticity can facilitate reproductive strategies that maximize mating success in variable environments and lead to differences in sex allocation among populations. For simultaneous hermaphrodites with sperm competition, including Serranus tortugarum a small coral reef fish, proportional male allocation (testis in total gonad) is often greater where local density or mating group size is higher. We tested whether S. tortugarum reduced male allocation when transplanted from a higher density site to a lower density site. After 4 months, transplants mirrored the sex-allocation patterns of the resident population on their new reef. Transplants had significantly lower male allocation than representatives from their source population, largely as a result of reduced testis mass relative to body size.