Pilar Castro-Díez

Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types.

Abstract The structural causes of variation in leaf mass per area, and of variations in leaf structure accounted for by leaf habit and life form, were explored in a set of laboratory-grown seedlings of 52 European woody species. The leaf traits analysed included density, thickness, saturated mass/dry mass, and leaf nitrogen per mass and per area. Other traits described the anatomy of leaves, most of them relating to the lamina (proportions of palisade and spongy parenchymata, epidermis, air space and sclerified tissues, expressed as volume per leaf area, and per-cell transversal areas of epidermis and parenchymata), and another referring to the mid rib (transversal section of sclerified tissues). Across the whole set of species leaf mass per area was correlated with leaf density but not with thickness, and this was confirmed by taxonomic relatedness tests. Denser leaves corresponded with greater proportion of sclerified tissues in the lamina, smaller cells and lower water and N contents, but no relation was found with the proportion of air space in the lamina. Taxonomic relatedness analysis statistically supported the negative association of leaf density with saturated to dry leaf mass ratio. Thicker leaves also exhibited greater volume per leaf area and greater individual cell area in each of the tissues, particularly parenchyma. Mean leaf mass per area and leaf thickness were lower in deciduous than in evergreen species, but no significant differences in leaf density, proportion of sclerified tissues in the lamina or cell area were found between the two groups. Leaf mass per area was higher in trees and subshrubs than in shrubs and climbers-plus-scramblers, this rank being equal for leaf density and proportion of sclerified tissues in the lamina, and reversed for cell area. Given the standardised environment and ontogenetic phase of the seedlings, we conclude that variation in leaf structure and anatomy among species and species groups has a strong genetic basis, and is already expressed early in the development of woody plants. From an ecological viewpoint, we can interpret greater leaf mass per area across this species set as greater allocation to support and defence functions, as shown predominantly by species from resource-poor environments.

Stem anatomy and relative growth rate in seedlings of a wide range of woody plant species and types

Abstract Stem traits were analysed in laboratory-grown seedlings of 80 European woody and semiwoody species of known potential relative growth rate (RGR) and of similar ontogenetic phase. The objectives were, firstly, to assess the relation between stem structure and plant growth potential and, secondly, to explore how stem structure varies among species differing in life form and leaf habit. Hydraulic conductance was represented by the mean diameter of the widest xylem conduits (Dmax), and structural strength by the percentage of xylem tissue occupied by cell wall material (CWx) or stem tissue density (SD). Across all species RGR showed a weak positive correlation with Dmax and weak negative ones with CWx and SD, with slow-growers showing great dispersion of stem trait values. In the RGR-Dmax relationship this dispersion disappeared when trees were removed from the analysis. None of the relationships were significant among tree species alone. It was suggested that fast-growers require a xylem with wide conduits (high Dmax) to achieve high hydraulic conductivity, and “cheaply” constructed stems (low CWx and SD) to maximise allocation to leaves. However, the possession of such traits does not guarantee fast growth, as other factors may constrain RGR elsewhere in the plant. Deciduous seedlings showed higher Dmax and lower CWx than evergreens. Higher Dmax could reflect an innate higher tolerance of conductivity loss by freeze-induced embolism in deciduous plants, which are not burdened by the maintenance of foliage in winter. In contrast, life forms were differentiated most clearly by SD. For instance, shrub seedlings had less dense stem tissues than tree seedlings, possibly because they need less investment in long-term strength and stature.

What explains variation in the impacts of exotic plant invasions on the nitrogen cycle? A meta‐analysis

Exotic plant invasions can notably alter the nitrogen (N) cycle of ecosystems. However, there is large variation in the magnitude and direction of their impact that remains unexplained. We present a structured meta-analysis of 100 papers, covering 113 invasive plant species with 345 cases of invasion across the globe and reporting impacts on N cycle-related metrics. We aim to explain heterogeneity of impacts by considering methodological aspects, properties of the invaded site and phylogenetic and functional characteristics of the invaders and the natives. Overall, plant invasions increased N pools and accelerated fluxes, even when excluding N-fixing invaders. The impact on N pools depended mainly on functional differences and was greater when the invasive plants and the natives differed in N-fixation ability, plant height and plant/leaf habit. Furthermore, the impact on N fluxes was related mainly to climate, being greater under warm and moist conditions. Our findings show that more functionally distant invaders occurring in mild climates are causing the strongest alterations to the N cycle.

Flowering phenology of invasive alien plant species compared with native species in three Mediterranean-type ecosystems

BACKGROUND AND AIMS Flowering phenology is a potentially important component of success of alien species, since elevated fecundity may enhance invasiveness. The flowering patterns of invasive alien plant species and related natives were studied in three regions with Mediterranean-type climate: California, Spain and South Africas Cape region. METHODS A total of 227 invasive-native pairs were compared for seven character types across the regions, with each pair selected on the basis that they shared the same habitat type within a region, had a common growth form and pollination type, and belonged to the same family or genus. KEY RESULTS Invasive alien plant species have different patterns of flowering phenology from native species in the three regions. Whether the alien species flower earlier, later or at the same time as natives depends on the climatic regime in the native range of the aliens and the proportion of species in the invasive floras originating from different regions. Species invading at least two of the regions displayed the same flowering pattern, showing that flowering phenology is a conservative trait. Invasive species with native ranges in temperate climates flower earlier than natives, those from Mediterranean-type climates at the same time, and species from tropical climates flower later. In California, where the proportion of invaders from the Mediterranean Basin is high, the flowering pattern did not differ between invasive and native species, whereas in Spain the high proportion of tropical species results in a later flowering than natives, and in the Cape region early flowering than natives was the result of a high proportion of temperate invaders. CONCLUSIONS Observed patterns are due to the human-induced sympatry of species with different evolutionary histories whose flowering phenology evolved under different climatic regimes. The severity of the main abiotic filters imposed by the invaded regions (e.g. summer drought) has not been strong enough (yet) to shift the flowering pattern of invasive species to correspond with that of native relatives. It does, however, determine the length of the flowering season and the type of habitat invaded by summer-flowering aliens. Results suggest different implications for impacts at evolutionary time scales among the three regions.

Trade-offs between phenology, relative growth rate, life form and seed mass among 22 Mediterranean woody species

Mediterranean woody plants exhibit a wide phenological diversity which cannot be explained just on the basis of climatic constraints. We assessed the role of relative growth rate (RGR), life form, seed and fruit mass as potential constraints of plant phenology. In a comparison of traits of 22 Mediterranean woody plant species, the duration of the primary shoot growing period decreased from climbers to shrubs and to trees. A hypothesised negative association between RGR and primary shoot growth duration did not emerge in our species set. The mechanism underlying phenological differences between plant life forms might relate to differences in the proportion of respiring to photosynthetic tissues, which decreases from climbers to trees. It is suggested that the degree of shoot preformation within the bud correlates with primary shoot growth duration, but not with RGR. Development of big fruits and seeds competes for carbon with vegetative growth. Indeed, species with bigger seeds and fruits exhibited shorter primary shoot growing periods, which tended to overlap with flower bud formation and flowering periods. We suggest that duration of primary shoot growth allow to short out the species between two extreme growth strategies: The conservative one would be characterised by a concentration of the primary shoot growth into a short period, free of frosts and droughts, and by a diversion of part of the current resources to assure next years growth. The opportunistic strategy, on the other extreme, would be defined by the allocation of resources to current growth whenever they are available, achieving longer growing periods at the expense of higher risk of tissue damage. These strategies should have been selected for in environments of predictable and unpredictable resource availability, respectively.

The relative importance for plant invasiveness of trait means, and their plasticity and integration in a multivariate framework.

Functional traits, their plasticity and their integration in a phenotype have profound impacts on plant performance. We developed structural equation models (SEMs) to evaluate their relative contribution to promote invasiveness in plants along resource gradients. We compared 20 invasive-native phylogenetically and ecologically related pairs. SEMs included one morphological (root-to-shoot ratio (R/S)) and one physiological (photosynthesis nitrogen-use efficiency (PNUE)) trait, their plasticities in response to nutrient and light variation, and phenotypic integration among 31 traits. Additionally, these components were related to two fitness estimators, biomass and survival. The relative contributions of traits, plasticity and integration were similar in invasive and native species. Trait means were more important than plasticity and integration for fitness. Invasive species showed higher fitness than natives because: they had lower R/S and higher PNUE values across gradients; their higher PNUE plasticity positively influenced biomass and thus survival; and they offset more the cases where plasticity and integration had a negative direct effect on fitness. Our results suggest that invasiveness is promoted by higher values in the fitness hierarchy--trait means are more important than trait plasticity, and plasticity is similar to integration--rather than by a specific combination of the three components of the functional strategy.

Different flowering phenology of alien invasive species in Spain: evidence for the use of an empty temporal niche?

Flowering phenology is an important and poorly understood plant trait that may possibly be related to the invasiveness potential of alien species. The present work evaluates whether flowering time of invasive alien species is a key trait to overcome the climatic filters operating in continental Mediterranean ecosystems of Spain (characterised by summer drought and low temperatures in winter). We conducted comparisons between the flowering phenology of the invasive species in their native range and in Spain, and between flowering phenology of 91 coexisting invasive-native species pairs. For the alien species, geographical change from the native to the invaded region did not result in shifts in the start and the length of the flowering period. Overall, climatic conditions in the native range of species selected for a flowering pattern is maintained after translocation of the species to another region. Flowering of tropical and temperate invasive alien species peaked in summer, which contrasts with the spring flowering of native and invasive alien species of Mediterranean climate origin. By exploiting this new temporal niche, these invasive alien species native to tropical and temperate regions benefit from reduced competition with natives for abiotic and biotic resources. We suggest that human-mediated actions have reduced the strength of the summer drought filter in particular microhabitats, permitting the invasion of many summer-flowering aliens.

Differential and interactive effects of temperature and photoperiod on budburst and carbon reserves in two co-occurring Mediterranean oaks

Effects of temperature and photoperiod and their interactions on budburst and on the use of carbon reserves were examined in two Mediterranean oaks differing in wood anatomy and leaf habit. Seedlings of Quercus ilex subsp. ballota (evergreen and diffuse-porous wood) and Q. faginea (semi-deciduous and ring-porous wood) were grown under two temperatures (12 and 19 degrees C) and two photoperiods (10 and 16 h) in a factorial experiment. In the 16 h photoperiod at 19 degrees C, photosynthesis was suppressed in half of the seedlings by covering leaves with aluminium foil. The concentration of soluble sugars, starch and lipids in leaves, stems and roots was assessed before and after budburst. Under the 12 degrees C treatment (mean current temperature in early spring in the Iberian Peninsula), budburst in Q. faginea occurred earlier than in Q. ilex. Higher temperature promoted earlier budburst in both species, mostly under the 16 h photoperiod. This response was less pronounced in Q. faginea because its budburst was also controlled by photoperiod, and because this species needs to construct a new ring of xylem before budburst to supply its growth demands. Therefore, dates of budburst of the two species became closer to each other in the warmer treatment, which might alter competitive relations between the species with changing climate. While Q. ilex relied on carbon reserves for budburst, Q. faginea relied on both carbon reserves and current photoassimilates. The different responses of the two Quercus species to temperature and photoperiod related more to xylem structure than to the source of carbon used for budburst.

Seasonal carbon storage and growth in Mediterranean tree seedlings under different water conditions

In all Mediterranean-type ecosystems, evergreen and deciduous trees differing in wood anatomy, growth pattern and leaf habit coexist, suggesting distinct adaptative responses to environmental constraints. This study examined the effects of summer water stress on carbon (C) storage and growth in seedlings of three coexisting Mediterranean trees that differed in phenology and wood anatomy characteristics: Quercus ilex subsp. ballota (Desf.) Samp., Quercus faginea Lam. and Pinus halepensis L. Seedlings were subjected to two levels of watering during two consecutive summers and achieved a minimum of -0.5 and -2.5 MPa of predawn water potential in the control and water stress treatment, respectively. Both Quercus species concentrated their growth in the early growing season, demanding higher C in early spring but replenishing C-stores in autumn. These species allocated more biomass to roots, having larger belowground starch and lipid reserves. Quercus species differed in seasonal storage dynamics from P. halepensis. This species allocated most of its C to aboveground growth, which occurred gradually during the growing season, leading to fewer C-reserves. Soluble sugar and starch concentrations sharply declined in August in P. halepensis, probably because reserves support respiration demands as this species closed stomata earlier under water stress. Drought reduced growth of the three species, mainly in Q. faginea and P. halepensis, but not C-reserves, suggesting that growth under water stress conditions is not limited by C-availability.

The exotic aquatic mud snail Potamopyrgus antipodarum (Hydrobiidae, Mollusca): state of the art of a worldwide invasion

Biological invasions represent a relevant ecological and economic problem of our globalized world. While a few species have been classified as invasive due to their ecological and economic impacts on the invaded ecosystems (e.g., zebra mussel), others show contrasting invasive potential, depending on the invaded ecosystem and/or the traits of the exotic species. This paper reviews the worldwide distribution, ecological impacts and the reasons that explain the invasive success of the aquatic mud snail Potamopyrgus antipodarum Gray (Hydrobiidae, Mollusca), which is native to New Zealand. This review shows that most studies on P. antipodarum distribution have been conducted in Europe, North America and Australia, and few studies in Asia. The distribution of this snail is still unknown in other parts of the world (e.g., Africa, South and Central America). The range of invaded aquatic ecosystems varies from fresh to salt water and from lentic to lotic ecosystems. The ecological impact of this species is due to the fast population growth rate and to the extremely high densities that it can reach, leading to altered C and N cycles in invaded ecosystems. However, at low densities mud snails have been shown to enhance secondary production. Additionally, P. antipodarum has been found to overcome the negative effects of predators and parasites (e.g., it survives the pass through the digestive tracts of fish). This review contributes to assess the magnitude and ecological risk of P. antipodarum invasion throughout the world.