How do evergreen and deciduous species respond to shade?—Tolerance and plasticity of subtropical tree and shrub species of South-East China

Different plant species make use of resource gradients such as light in different ways. First, plant species specialize in using different parts of the gradient, resulting in niche partitioning, Second, within the section of the gradient used by a species, plants respond to a different resource supply by plasticity. Our study addressed both of these strategies, with the main objective to relate mean responses and plasticity indices of seedlings of woody species to species characteristics such as leaf habit and to variation in branch lengths and local frequency of adult trees in forest communities.A greenhouse experiment with 36 deciduous and 35 evergreen subtropical tree and shrub species was carried out to test the influence of light and nutrient availability on trait expression and plasticity of the species. The greenhouse responses of seedlings were compared with adult individuals in the field, based on a set of 46 species that occurred also in the 27 permanent plots in a secondary subtropical broadleaved forest in Zhejiang Province (SE-China).In the greenhouse experiment, most variables showed significant differences between unshaded (250 μE m⁻² s⁻¹) and shaded (10 μE m⁻² s⁻¹) treatments as well as between high and low nutrient supply. Deciduous species were more plastic than evergreen species in their response to light. No significant correlations were detected in mean responses and plasticity between juvenile plants in the greenhouse and adult trees in the field. However, some trait values such as biomass and node density as well as plasticity of several traits were positively related to the species’ abundance in the field, suggesting that locally abundant species tend to be less “plastic” than locally rare species.     

Chemical and morphological variation of Mediterranean woody evergreen species: Do plants respond to ungulate browsing?

Abstract. Plant defense theories suggest that chemical or structural defences should be maximized when and where browsing is most likely to occur. We tested this hypothesis on four evergreen woody species growing in a Mediterranean area with a high density of ungulates. In this system, levels of browsing are more intense in the winter (due to the lack of annual plants) and young foliage is often preferred. Therefore we predicted that the chemical defences of these species, namely their phenolic content, would vary with leaf age, season and damage intensity. In addition, we tested whether ungulates preferentially selected species containing lower phenolic levels, and also whether browsing induced either chemical or morphological changes in damaged plants. Phenolic levels varied greatly between plant species; ungulates browsed preferentially on the species with the lowest phenolic levels. No difference in phenolic content was found between browsed and unbrowsed trees. Morphological changes in heavily browsed trees included an increase in shoot and leaf density and a net decrease in leaf size. We suggest that for Mediterranean plants, which have evolved under high browsing pressure from large mammals, the production of small leaves and dense shoots in response to browsing might decrease ungulate foraging efficiency and hence reduce the rate of further damage as effectively as high levels of chemical defence.     

Habitat selection and ranges of tolerance: how do species differ beyond critical thresholds?

Sensitivity to habitat fragmentation often has been examined in terms of thresholds in landscape composition at which a species is likely to occur. Observed thresholds often have been low or absent, however, leaving much unexplained about habitat selection beyond initial thresholds of occurrence, even for species with strong habitat preferences. We examined responses to varying amounts of tree cover, a widely influential measure of habitat loss, for 40 woodland bird species in a mixed woodland/grassland landscape in eastern North Dakota, USA. We used LOESS smoothing to describe incidence for each species at three scales: within 200, 400, and 1200 m around sample locations. For the 200‐m scale, we also calculated the most‐preferred range of tree cover (within which at least half of observations were predicted to occur) for each species. Only 10 of 40 species had occurrence thresholds greater than about 10% tree cover. After initial occurrence, species showed three general patterns: some increased monotonically with tree cover; some increased up to an asymptote; some peaked at intermediate amounts of tree cover and then declined. These patterns approximate selection for interior woodlands and for edge‐rich environments, but incidence plots provide greater detail in landscape‐scale selection than do those categories. For most species, patterns persisted at larger scales, but for some, larger scales had distinctly different patterns than local scales. Preferred ranges of tree cover varied from <20% tree cover (common grackle, Quiscalus quiscula) to >60% (veery, Catharus fuscescens). We conclude that incidence patterns provide more information on habitat selection than do threshold measures for most species: in particular, they differentiate species preferring concentrated woodlands from those preferring mixed landscapes, and they show contrasting degrees of selectiveness. [Correction added on 16 October 2012, after first online publication: the Abstract section has been reworded].     

Functional traits of woody plants: correspondence of species rankings between field adults and laboratory‐grown seedlings?

Abstract. Research into interspecific variation in functional traits is important for our understanding of trade‐offs in plant design and function, for plant functional type classifications and for understanding ecosystem responses to shifts in species composition. Interspecific rankings of functional traits are a function of, among other factors, ontogenetic or allometric development and environmental effects on phenotypes. For woody plants, which attain large size and long lives, these factors might have strong effects on interspecific trait rankings. This paper is the first to test and compare the correspondence of interspecific rankings between laboratory grown seedlings and field grown adult plants for a wide range of functional leaf and stem traits. It employs data for 90 diverse woody and semiwoody species in a temperate British and a (sub)Mediterranean Spanish flora, all collected according to a strict protocol. For 12 out of 14 leaf and stem traits we found significant correlations between the species ranking in laboratory seedlings and field adults. For leaf size and maximum stem vessel diameter > 50 % of variation in field adults was explained by that in laboratory seedlings. Two important determinants of plant and ecosystem functioning, specific leaf area and leaf N content, had only 27 to 36 and 17 to 31 % of variation, respectively, in field adults explained by laboratory seedlings, owing to subsets of species with particular ecologies deviating from the general trend. In contrast, interspecific rankings for the same traits were strongly correlated between populations of field adults on different geological substrata. Extrapolation of interspecific trait rankings from laboratory seedlings to adult plants in the field, or vice versa, should be done with great caution.     

Living on the edge: do central and marginal populations of plants differ in habitat suitability?

The performance of populations at the edge of specie’s distribution range may differ substantially from central populations. Here, we develop a modelling framework to estimate ecological niches (i.e. climatic) of four locally endangered plant species and measure the distance of marginal (geographically) populations to the species’ niche centroid in order to analyse whether marginal populations are outside of the optimal ecological niche of each species. Our results show that for three of the four studied species, which have their populations located at the margins of their distribution ranges, are also at the margins of their climate gradients. These results would support the hypothesis that marginality within the set of habitable conditions (i.e. ecological niche) may represent an important determinant on performance of some plant populations, as well as a plausible explanation to the degree of stagnancy or regression experienced by species in those regions where their populations are at the margin of their ecological niche and/or they are restricted to microrefugia with ecological conditions very different from those around them. Finally, this study aims to be a theoretical base from which to advance on, including other types of factors (e.g. biotic interactions, topography, human influence and population fluctuations through time), which will allow for a better understanding of the complex network of factors that occur in marginal plant populations.     

Combined effects of shade and drought on tulip poplar seedlings: trade‐off in tolerance or facilitation?

It has been hypothesised that many natural plant distribution patterns can be explained by a trade-off between shade and drought tolerance which would make plants more sensitive to shade under dry conditions. On the other hand, shading by nurse plants is often observed to enhance growth and survival of smaller plants in dry areas. This paper describes an experiment designed to address the interactive effects of drought and shade on growth and physiology of tulip poplar seedlings ( Liriodendron tulipifera ) grown in shade houses under different combinations of irradiance (1%, 5%, 12%, 17%, and 27% of ambient PAR) and soil water content (5–9%, 11–15%, and>20%). The results show no evidence that higher PAR levels compensate for the negative effects of drought on photosynthesis and growth. Rather, rates of estimated daily shoot carbon gain decreased with PAR under dry conditions. Daily xylem sapflow patterns indicated that this was associated with a strong reduction in stomatal conductance in plants growing in dry soil and high PAR conditions. Whole-shoot light compensation points were higher for plants raised under higher PAR conditions, but were not significantly influenced by the water treatments. Shoot dark respiration rates decreased under drier conditions. These results do not support the hypothesis of a trade-off between shade and drought tolerance. Instead, they indicate mechanisms that can lead to positive effects of shading by neighbouring plants under dry conditions. Indeed, such facilitation is thought to be important in many plant communities.     

Does shade improve light interception efficiency? A comparison among seedlings from shade-tolerant and -intolerant temperate deciduous tree species

Here, we tested two hypotheses: shading increases light interception efficiency (LIE) of broadleaved tree seedlings, and shade-tolerant species exhibit larger LIEs than do shade-intolerant ones. The impact of seedling size was taken into account to detect potential size-independent effects on LIE. LIE was defined as the ratio of mean light intercepted by leaves to light intercepted by a horizontal surface of equal area. Seedlings from five species differing in shade tolerance (Acer saccharum, Betula alleghaniensis, A. pseudoplatanus, B. pendula, Fagus sylvatica) were grown under neutral shading nets providing 36, 16 and 4% of external irradiance. Seedlings (1- and 2-year-old) were three-dimensionally digitized, allowing calculation of LIE. Shading induced dramatic reduction in total leaf area, which was lowest in shade-tolerant species in all irradiance regimes. Irradiance reduced LIE through increasing leaf overlap with increasing leaf area. There was very little evidence of significant size-independent plasticity of LIE. No relationship was found between the known shade tolerance of species and LIE at equivalent size and irradiance.     

Does irrigation affect leaf phenology in deciduous and evergreen trees of the savannas of northern Australia?

Abstract Soil moisture was augmented experimentally during two successive dry seasons and the intervening wet season in a humid tropical savanna in Darwin, northern Australia. Leaf phenology was monitored in four common tree species Termmalia ferdinandiana and Planchonia careya (both deciduous species), and Eucalyptus miniata and Eucalyptus tetrodonta (both evergreen species). Irrigation produced consistently significant effects in only T. ferdinandiana. In this species leaf-flush was significantly earlier, canopy decline and leaf-fall were significantly later and the attainment of full canopy was earlier in irrigated compared with non-irrigated trees. Litterfall, and the seasonal patterns of contraction and expansion of stems (a measure of stem water status or storage) were not significantly affected by irrigation in any species. Leaf longevity in the deciduous species was 6–8 months; some eucalypt leaves lived for approximately 1 year, but none lived longer than 18 months. Irrigation had relatively little effect on leaf longevity. While variation in soil moisture is a potentially important cue to both leaf-fall and leaf-flush, stem water status and climatic factors such as vapour pressure deficit may also be important climatic cues to phenological behaviour.     

From evergreen to deciduous tropical forests: how energy–water balance,temperature, and space influence the tree species composition in a high diversity region

Background: Understanding floristic and geographic patterns in one of the most biodiverse regions in the world – the Atlantic forest of eastern Bahia, Brazil – can identify the drivers of diversity in tropical forests and provide useful information for biological conservation.

Aims: To understand the role of both climate and geographical location on variation in tree species composition in a region characterised by an abrupt transition from wet forests to semi-arid thorn-woodlands. To test whether a regional classification of forests according to elevation belts and leaf flush pattern is consistent with floristic composition.

Methods: We submitted 14,094 tree species occurrence records and 31 geo-climatic variables prepared for 57 sites in eastern Bahia, Brazil, to multivariate and regression analyses and variance partitioning.

Results: Climate and space were both significantly (P ≤ 0.05) contributing to explaining floristic variations. Actual evapotranspiration, duration of water deficit, and minimum temperature of coldest month were the main predictors. Floristic differences were significant except when comparing evergreen lower plains and upper plains forests.

Conclusions: Although distance among sites may play an important role, species composition is chiefly influenced by environmental gradients. This highlights environmental heterogeneity as a key factor in the planning of biodiversity conservation in tropical forests.     

What do we know about growth of vessel elements of secondary xylem in woody plants?

Despite extensive knowledge about vessel element growth and the determination of the axial course of vessels, these processes are still not fully understood. They are usually explained as resulting primarily from hormonal regulation in stems. This review focuses on an increasingly discussed aspect – mechanical conditions in the vascular cambium. Mechanical conditions in cambial tissue are important for the growth of vessel elements, as well as other cambial derivatives. In relation to the type of stress acting on cambial cells (compressive versus tensile stress) we: (i) discuss the shape of the enlarging vessel elements observed in anatomical sections; (ii) present hypotheses regarding the location of intrusive growth of vessel elements and cambial initials; (iii) explain the relationship between the growth of vessel elements and fibres; and (iv) consider the effect of mechanical stress in determining the course of a vessel. We also highlight the relationship between mechanical stress and transport of the most extensively studied plant hormone – auxin. We conclude that the integration of a biomechanical factor with the commonly acknowledged hormonal regulation could significantly enhance the analysis of the formation of vessel elements as well as entire vessels, which transport water and minerals in numerous plant species.     

Peptidomics of neurohemal organs from species of the cockroach family Blattidae: how do neuropeptides of closely related species differ?

The primary structures and molecular mass data of neuropeptides that are released from major neuroendocrine sites of the American cockroach, Periplaneta americana, are well known. In the current study we extensively surveyed neuropeptides of the corpora cardiaca, the corpora allata and the abdominal perisympathetic organs, from 14 related cockroach species belonging to the family Blattidae. Mainly, this survey was executed by using mass spectrometric methods (MALDI-TOF MS). Peptides which appeared to be modified, as judged from their mass data and comparison with known data from P. americana, were fragmented by means of the post-source decay technique to determine their amino acid sequence. Single organ preparations sufficed to reveal the peptide pattern in neurohemal organs and to identify species-specific modifications, making it possible to glean information also from tissues of less abundant species that were caught during field trips. The peptide inventory described in this study is very typical of cockroaches of the family Blattidae and clearly separates this taxon from other taxa of cockroaches. The majority of neuropeptides was identical in all investigated species. Some peptides, however, displayed remarkable variations in their sequence which hint to a differential rate of modification among peptides of the same neuropeptide family. These modifications serve to distinguish further relatedness of taxa within the Blattidae.     

Do species of invaded communities differ in their vulnerability to being eliminated by the dominant alien plants?

Invasions are associated with loss of diversity and changes in species composition. This study aims to provide evidence if this loss happens at random or if the loss is related to functional traits of native species. Traits of species in communities dominated by 13 plants alien to Central Europe were compared to those found in the adjacent uninvaded vegetation. There is a weak but significant non-random pattern in the distribution of traits between the invaded and adjacent uninvaded vegetation. For example, species possessing a taproot, annuals but also juveniles of trees tend to be proportionally more abundant in the invaded vegetation, suggesting they are impacted less than species with high clonal index, perennial polycarpic species and species without a taproot, which are proportionally more represented in the uninvaded vegetation. Fast laterally spreading species, trees’ juveniles and woody species in general are proportionally more frequent in the invaded vegetation, while biennials, slow laterally spreading species but also shrubs are proportionally more frequent in the uninvaded vegetation. Juveniles of trees may compete successfully with the aliens due to being adapted to thrive in low light conditions. Annuals may thrive in the invaded communities by possessing a life strategy different from most of the selected aliens. Possessing a taproot appears to be another trait important for the successful coexistence with the dominant invasive aliens, possibly pointing to the importance of underground competition. Clonal perennial polycarpic herbs are species functionally most similar to most of the selected aliens, and these species were found to be most underrepresented in the invaded vegetation.     

Few immigrant phytophagous insects on woody plants in Europe: legacy of the European crucible?

Exotic phytophagous insects are invading forest ecosystems worldwide. So far, 109 invasive insects on woody plants, 57 from North American (NA), and 52 from Asia (A) have established populations in European forests. Four orders account for about 84% of the immigrants: Homoptera 39%, Lepidoptera 13%, Coleoptera 19%, and Hymenoptera 13%. The majority of these invasive species (63% of NA and 77% of A) live on deciduous trees, of which 36% have been introduced from NA and Asia. The remaining insect species (37% NA and 25% A) live on various conifers, of which 53% have also been introduced. Most (57%) of the NA insects feeding on coniferous plants live upon their introduced, native host plants. These data suggest that many NA immigrant phytophagous species in Europe have been successful in establishing permanent populations because their native hosts preceded or accompanied them into Europe and/or were asexually reproducing species. We propose that fewer invasive phytophagous insects have become established in European compared to North American woodlands because of the unique legacy of the European Pleistocene/Holocene crucible (i.e. endless cycles of populations contracting into highly disparate, dispersed metapopulation refugia and eventually expanding out of them) on European species and ecosystems that caused highly diminished heterogeneity. This translates to fewer and less penetrable tri-trophic niches in Europe due to fewer and less available host plants, but greater zootic resistance per niche derived from more competition-hardened competitors and possibly natural enemies. Moreover, many European species are probably superior invasion specialists because the crucible favored traits that are conducive to success in highly subdivided, and extinction-prone metapopulations: asexual reproduction, polyploidy, and other traits especially conducive to persistence under stress, and explosive growth/spread under amelioration.     

Litter decomposition of woody species in shrublands of NW Patagonia: how much do functional groups and microsite conditions influence decomposition?

The study examined the effects of leaf traits, soil microsite, and microclimate characteristics on litter decomposition of the dominant species in two functional groups (FG), deciduous and evergreen, in shrublands in NW Patagonia, Argentina. Leaf traits considered were nutrient concentration (C, N, P, C/N, and N/P) and physical characteristics (area, strength, specific leaf area, and dry matter content). Soil microsite characteristics measured were pH, C, N, P, C/N and water retention capacity, while soil microclimate characteristics recorded were soil and air, temperature and moisture, and solar radiation. Five evergreen and five deciduous woody shrub species were selected. During 1 year, litter and microsite properties were measured below canopy: (i) senescent leaf chemical and physical properties, and the quantity as well as field decomposition of litter and (ii) soil chemistry, and soil and air physical properties. The factors controlling litter decomposition were different for each FG. In deciduous species, C/N ratio had a negative effect on decomposition. In evergreen species, decomposition was affected negatively by leaf carbon and dry matter content. Litter decomposition depended exclusively on the inherent senescent leaves traits. The common decomposition pattern between species of both FG could be attributed to similar leaf traits and the correlation between variables that control decomposition in both groups. Plant nutrient inputs associated with the litter decomposition process did not explain the soil nutrient content. These results suggest that other organic matter sources (roots, branches, and fruits) are more important than leaves on soil fertility.     

The general occurrence of “plastid initials” and their development into plastids in cortical cells of woody plants in winter

Summary Electron microscopic studies revealed that plastid initials, presumed precursors of plastids, occur in cortical cells of the following plants studied in February and March: Betula ermanii Cham.; Prunus sargentii Rhed.; Pyrus communis L.; Ribes sinanense F. Maekawa; Salix matsudana Koidz. forma tortuosa Rhed.; and Sambucus sieboldiana var. miquelii Hara. Since plastid initials were found previously in Malus pumila Mill., Morus bombycis Koidz. and Populus euramericana cv. gelrica (Sagisaka 1991), plastid initials have been found in all woody plants examined to date. In P. euramericana cv. gelrica, at later stages of the development of the initials in March, the conglomerates of plastid initials became heterogeneous in terms of size, extent of thylakoid formation and ability to form starch granules. The formation of prolamellar structures was frequently observed in cells of Magnolia kobus var. borealis Sarg., which was sampled on April 19. These observations suggest the course of events in the development of the plastid initial and the continuity of the life of amyloplasts over a year in the life of woody plants.     

Allocation,stress tolerance and carbon transport in plants: how does phloem physiology affect plant ecology?

Despite the crucial role of carbon transport in whole plant physiology and its impact on plant–environment interactions and ecosystem function, relatively little research has tried to examine how phloem physiology impacts plant ecology. In this review, we highlight several areas of active research where inquiry into phloem physiology has increased our understanding of whole plant function and ecological processes. We consider how xylem–phloem interactions impact plant drought tolerance and reproduction, how phloem transport influences carbon allocation in trees and carbon cycling in ecosystems and how phloem function mediates plant relations with insects, pests, microbes and symbiotes. We argue that in spite of challenges that exist in studying phloem physiology, it is critical that we consider the role of this dynamic vascular system when examining the relationship between plants and their biotic and abiotic environment.