Emergence and survival of herbaceous seedlings in temperate grassy woodlands: Recruitment limitations and regeneration niche

Abstract Field experiments examined herbaceous seedling emergence and survival in temperate grassy woodlands on the New England Tablelands of New South Wales. Effects of intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds, grazing and seed theft by ants on seedling emergence and survival were studied in two field experiments. Thirteen species with a range of traits were used in the experiments and their cumulative emergence was compared with laboratory germination studies. Field emergence correlated to laboratory germination but all species had lower emergence in the field. Little natural emergence of native species was observed in the field in unsown treatments. Short‐lived forbs had the highest emergence, followed by perennial grasses; rhizomatous graminoids and perennial forbs had the lowest emergence. Soil surface and cover treatments did not markedly enhance emergence suggesting that intertussock spaces were not prerequisites for forb emergence. No consistent pattern of enhanced emergence was found for any treatment combination across all species. Seedling survival varied among species, with perennial grasses and short‐lived forbs having the highest seedling mortality. Low mortality rates in the graminoids and rhizomatous forbs appeared partially to compensate for lower seedling emergence. All perennial grasses and some short‐lived forbs showed increased risk of mortality with grazing. Differences in emergence and survival of species were related to ground cover heterogeneity, soil surfaces and, to some extent, herbivory. The complexity of these patterns when superimposed on temporal variability suggests that no generalizations can be made about the regeneration niche of herbaceous species groups. Strong recruitment limitation and partitioning of resources in the regeneration niche may reduce competition among native species and explain the high species richness of the herbaceous layer in the temperate grassy communities of eastern Australia.     

Effects of plant traits and the relative abundance of common woody species on seedling herbivory in the Thousand Island Lake region

Aims Plant-herbivore interaction is a hot topic in the study of biodiversity and ecosystem functions. Herbivores can negatively affect seedling growth and therefore can alter the dynamics of plant recruitment. However, previous studies do not fully reveal the relative importance of different plant functional traits on herbivory intensity and rarely link herbivory to the relative abundance of plant species.Methods Here, we measured 11 plant functional traits and the relative abundance of seedlings of 16 common woody species in the subtropical forests on 29 islands in Thousand Island Lake, East China. We then used multivariate regression and variance partitioning to test the contribution of functional traits and the relative abundance to interspecific differences of insect herbivory intensity.Important findings Our study found that both plant functional traits (e.g. carbon nitrogen ratio, leaf thickness) and the relative abundance of woody species played important roles in herbivory intensity, and they jointly contributed 54% of the variance of the interspecific differences. Among these factors, species with higher defensive ability, lower nutrient content and higher relative abundance had lower herbivory intensity. We suggest to consider both individual level traits (functional traits) and community level attributes (the relative abundance) in future herbivory studies.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Adaptations to water gradient in three Rorippa plant species correspond with plant resistance against insect herbivory under drought and waterlogged conditions

1. Plants live in environments where they are constantly, and often simultaneously, exposed to different types of biotic and abiotic stress, such as insect herbivory and water availability. How plants are adapted to abiotic conditions may determine how a surplus or shortage of water affects plant resistance to insect herbivory. Moreover, this effect may vary depending on the feeding mode of the herbivore.
2. We explored how three closely related Rorippa plant species that vary in adaptations to different water levels, resist herbivory by four different insects (aphids: Myzus persicae, Lipaphis erysimi, and caterpillars: Pieris brassicae, Plutella xylostella) under waterlogging or drought conditions. We hypothesized that plants that are differently adapted to water availability will be disparately affected by water availability in their resistance to insect herbivory.
3. On the semi-aquatic plant species Rorippa amphibia, both aphid species reached a larger colony size under drought conditions. This indicates that R. amphibia was compromised in resistance to aphid feeding when under drought conditions, to which it is less well adapted. Water conditions did not affect aphid performance on the flood-plain species Rorippa palustris. On the terrestrial plant species Rorippa sylvestris, aphids performed worse on waterlogged than drought-treated plants. Neither caterpillar species was significantly affected by the water availability of their food plant.
4. Our findings suggest that water availability can have distinct effects on plant–insect interactions. We propose that plant adaptations to water conditions can be a major predictor towards explaining the variation of effects that water availability can have on plant–insect interactions.

     

How species traits and affinity to urban land use control large-scale species frequency

Aim Although urban areas only occupy c. 2.8% of the earth's land surface, urbanization threatens biodiversity as areas of high human population density often coincide with high biodiversity. Therefore, nature conservation should concentrate on both remote areas and densely populated regions. Protecting rare plant species in rural and urban areas can contribute to the protection of biodiversity. We therefore need to understand why species are rare. Studies on causes of rarity often concentrate on either plant traits or extrinsic threats (such as habitat fragmentation or nitrogen enrichment). However, there are only a few studies that combine traits and extrinsic threats, although such analyses might clarify causes of rarity. We assessed how the affinity of vascular plant species to urban land use (‘urbanity’) interacts with plant traits in determining species frequency. Location Germany, resolution c. 12 km × 11 km. Methods Species with a low frequency may be rare because they occur in rare habitats or because of other reasons, although their habitat is frequent. Therefore, we calculated the frequency of species corrected for habitat frequency, i.e. relative species frequency. We explained relative species frequency by the interactions of species traits and species affinity to urban land use using generalized linear models. Simultaneous autoregressive error models controlled for phylogenetic relationships of species. Results Relative species frequency depends on species affinity to urban land use, independent of the different interactions between traits and urbanity used as predictors. The higher the urbanity the higher is species frequency. Urbanity interacts with species preferences towards temperature and soil acidity. Moreover, dispersal, nitrogen preferences and origin explain relative species frequency, amongst others. Main conclusions Many rare species, especially those preferring cool or acidic habitats might already have disappeared from urban areas. Analyses that combine species traits and environmental effects can explain the causes of rarity and help to derive better conservation strategies.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Untangling the relationships among regional occupancy,species traits,and niche characteristics in stream invertebrates

The regional occupancy and local abundance of species are affected by various species traits, but their relative effects are poorly understood. We studied the relationships between species traits and occupancy (i.e., proportion of sites occupied) or abundance (i.e., mean local abundance at occupied sites) of stream invertebrates using small‐grained data (i.e., local stream sites) across a large spatial extent (i.e., three drainage basins). We found a significant, yet rather weak, linear relationship between occupancy and abundance. However, occupancy was strongly related to niche position (NP), but it showed a weaker relationship with niche breadth (NB). Abundance was at best weakly related to these explanatory niche‐based variables. Biological traits, including feeding modes, habit traits, dispersal modes and body size classes, were generally less important in accounting for variation in occupancy and abundance. Our findings showed that the regional occupancy of stream invertebrate species is mostly related to niche characteristics, in particular, NP. However, the effects of NB on occupancy were affected by the measure itself. We conclude that niche characteristics determine the regional occupancy of species at relatively large spatial extents, suggesting that species distributions are determined by environmental variation among sites.     

Natural variation in life history strategy of Arabidopsis thaliana determines stress responses to drought and insects of different feeding guilds

Plants are sessile organisms and, consequently, are exposed to a plethora of stresses in their local habitat. As a result, different populations of a species are subject to different selection pressures leading to adaptation to local conditions and intraspecific divergence. The annual brassicaceous plant Arabidopsis thaliana is an attractive model for ecologists and evolutionary biologists due to the availability of a large collection of resequenced natural accessions. Accessions of A. thaliana display one of two different life cycle strategies: summer and winter annuals. We exposed a collection of 308 European Arabidopsis accessions, that have been genotyped for 250K SNPs, to a range of stresses: one abiotic stress (drought), four biotic stresses (Pieris rapae caterpillars, Plutella xylostella caterpillars, Frankliniella occidentalis thrips and Myzus persicae aphids) and two combined stresses (drought plus P. rapae and Botrytis cinerea fungus plus P. rapae). We identified heritable genetic variation for responses to the different stresses, estimated by narrow‐sense heritability. We found that accessions displaying different life cycle strategies differ in their response to stresses. Winter annuals are more resistant to drought, aphids and thrips and summer annuals are more resistant to P. rapae and P. xylostella caterpillars. Summer annuals are also more resistant to the combined stresses of drought plus P. rapae and infection by the fungus Botryris cinerea plus herbivory by P. rapae. Adaptation to drought displayed a longitudinal gradient. Finally, trade‐offs were recorded between the response to drought and responses to herbivory by caterpillars of the specialist herbivore P. rapae.     

Patterns of ramet population of Iris japonica Thunb. and their effects on herb diversity in different microsites on Jinyun Mountain, China

Spatial patterns of ramet population of Iris japonica Thunb. and their effect on species diversity in the herb layer of 3 microsites (open area of forest edge (OAFE), bamboo forest (BF) and evergreen broad-leaved forest (EBF)) on Jinyun Mountain were studied using spatial pattern, niche and diversity analyses in a combination of population and community methods. The results were as follows: (1) judged by V/m and Morisita index (Iδ), ramet population of I. japonica in 3 microsites all clumped from scale 0.5 m × 0.5 m to 2 m × 2 m; (2) the pattern scale and pattern intensity both gradually decreased on all scales, and the density of ramet population of I. japonica decreased with the increase in canopy density and the decrease in relative photon flux density (RPFD) and R/FR from OAFE to EBF. In OAFE and BF, widespread I. japonica had significantly negative influence on the dominance of original dominant species and on species diversity in the herb layer (p < 0.05), while those influences in EBF were extremely weak. The mechanisms that pattern characteristics of ramet population of I. japonica influence herb diversity in 3 microsites were different. In OAFE, strong regeneration niche (above-ground spatial and below-ground root) and trophic niche (nutrient) competition had significantly negative influence on the species diversity of rare herbs and dwarf herbs. In BF, strong regeneration niche (below-ground root) and trophic niche (above-ground for light and below-ground for nutrient) competition had negative effect on the occurrence of rare species and on the survival of other herb species. In EBF, weak niche competition had little effect on the survival of herb species. Intensity of regeneration niche and trophic niche competition between I. japonica and other herb species is the determinant to the mechanism that ramet population of I. japonica influences herb diversity.     

Rainfall seasonality and drought performance shape the distribution of tropical tree species in Ghana

Tree species distribution in lowland tropical forests is strongly associated with rainfall amount and distribution. Not only plant water availability, but also irradiance, soil fertility, and pest pressure covary along rainfall gradients. To assess the role of water availability in shaping species distribution, we carried out a reciprocal transplanting experiment in gaps in a dry and a wet forest site in Ghana, using 2,670 seedlings of 23 tree species belonging to three contrasting rainfall distributions groups (dry species, ubiquitous species, and wet species). We evaluated seasonal patterns in climatic conditions, seedling physiology and performance (survival and growth) over a 2‐year period and related seedling performance to species distribution along Ghana's rainfall gradient. The dry forest site had, compared to the wet forest, higher irradiance, and soil nutrient availability and experienced stronger atmospheric drought (2.0 vs. 0.6 kPa vapor pressure deficit) and reduced soil water potential (?5.0 vs. ?0.6 MPa soil water potential) during the dry season. In both forests, dry species showed significantly higher stomatal conductance and lower leaf water potential, than wet species, and in the dry forest, dry species also realized higher drought survival and growth rate than wet species. Dry species are therefore more drought tolerant, and unlike the wet forest species, they achieve a home advantage. Species drought performance in the dry forest relative to the wet forest significantly predicted species position on the rainfall gradient in Ghana, indicating that the ability to grow and survive better in dry forests and during dry seasons may allow species to occur in low rainfall areas. Drought is therefore an important environmental filter that influences forest composition and dynamics. Currently, many tropical forests experience increase in frequency and intensity of droughts, and our results suggest that this may lead to reduction in tree productivity and shifts in species distribution.     

Callus initiation and regeneration capacities in Brassica species

In order to know the genetic differences of de- and redifferentiation capacities, seven Brassica species (B. campestris, B. nigra, B. oleracea, B. hirta, B. carinata, B. juncea and B. napus) were cultured in vitro, and their response to the medium supplemented with various combinations of auxin and cytokinin hormones was compared. Important factors for callus initiation were shown to be auxin and species. Calli were induced most frequently in Murashige and Skoog (MS) medium with 1.0 mg/l 2,4-dichlorophenoxyacetic acid (2,4-D), whereas -naphthaleneacetic acid (NAA) induced preferentially roots at a concentration of 2 to 5 mg/l. Callus-, root- and shoot-forming capacities from explanted cotyledon tissues were significantly different among the seven Brassica species. Calli derived from cotyledons and hypocotyls of seven species were transferred to MS media with 20g/l sucrose, 0 to 0.1 mg/l NAA and 0 to 4 mg/l kinetin to compare their regeneration capacities. Among the seven species tested, B. napus (2n=4x=38, genome AACC) had the highest shoot forming capacity (20.0%). Other amphiploid species, B. carinata (2n=4x=34, BBCC) and B. juncea (2n=4x=36, AABB) formed shoots at low frequencies (2.8% and 1.2%, respectively). A diploid species, B. oleracea (2n=2x 18, CC) also showed high shoot formation (10.2% on average). This result suggests that the gene(s) controlling shoot formation may be localized in the C genome. Differences were also found among varieties and cultivars within a species. One of the cultivars, Siberian kale (B. oleracea var. acephala) gave about 50% shoot formation. This kale was shown cytologically to be an autotetraploid (2n=4x=36, CCCC), thus probably possessing a double set of the shoot-forming gene(s).     

Nurse species and indirect facilitation through grazing drive plant community functional traits in tropical alpine peatlands

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant–plant–herbivore interactions in stressful environments. We set up a 2‐year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion‐forming plants (hereafter referred to as “nurse” species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the “grazer–nurse–beneficiary” interaction module can be more complex than expected when evaluated in the field.     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.     

我国亚热带典型树种抗火性状随林龄和器官的变异

在湖南省炎陵县青石冈国有林场选择杨梅、木荷、山矾、润楠、甜槠、青冈6种亚热带典型树种,通过测定植物不同龄组(幼龄林、中龄林、成熟林)和器官(叶、枝、皮)等指示抗火性状的形态及物理和化学性质,探究同一树种不同器官、龄组间抗火性差异以及不同树种抗火性的综合排序。结果表明: 抗火性在不同器官和龄组间均呈现显著差异;相对于枝和皮,叶片具有较高的含水率(53.7%)、较高的粗灰分含量(4.5%)和较低的粗纤维含量(23.9%);随着林龄的增长,树木抗火性呈现先降后增的规律,中龄林的含水率、粗灰分和粗纤维含量最低。各器官的综合抗火性能在不同物种间差异显著。树叶的抗火性排序为:润楠>甜槠>山矾>杨梅>木荷>青冈;树枝的综合抗火性表现为润楠、甜槠最强,其次是杨梅、木荷;树皮表现为木荷和甜槠的抗火性较强,抗火性最弱的为润楠和青冈。各物种的综合抗火性能综合得分呈现分异特征,其中抗火性最强的为木荷(1.033)和杨梅(0.526),润楠(-0.405)和青冈(-1.151)的抗火性最弱。因此,木荷和杨梅是我国南方亚热带森林防火林带的首选树种。     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.