Dioecy and its correlates in the flowering plants

Considerable effort has been spent documenting correlations between dioecy and various ecological and morphological traits for the purpose of testing hypotheses about conditions that favor dioecy. The data analyzed in these studies, with few exceptions, come from local floras, within which it was possible to contrast the subsets of dioecious and nondioecious taxa with regard to the traits in question. However, if there is a strong phylogenetic component to the presence or absence of dioecy, regional sampling may result in spurious associations. Here, we report results of a categorical multivariate analysis of the strengths of various associations of dioecy with other traits over all flowering plants. Families were scored for presence of absence of monoecy or dioecy, systematic position, numbers of species and genera, growth forms, modes of pollination and dispersal, geographic distribution, and trophic status. Seven percent of angiosperm genera (959 of 13,500) contain at least some dioecious species, and ≈6% of angiosperm species (14,620 of 240,000) are dioecious. The most consistent associations in the data set relate the presence of dioecy to monoecy, wind or water pollination, and climbing growth. At both the family and the genus level, insect pollination is underrepresented among dioecious plants. At the family level, a positive correlation between dioecy and woody growth results primarily from the association between dioecy and climbing growth (whether woody or herbaceous) because neither the tree nor the shrub growth forms alone are consistently correlated with a family's tendency to include dioecious members. Dioecy appears to have evolved most frequently via monoecy, perhaps through divergent adjustments of floral sex ratios between individual plants. Monoecy itself is related to abiotic pollination and climbing growth as revealed by multivariate analysis. Dioecy and monoecy are concentrated in the less advanced superorders of Thorne (1992) and subclasses of Cronquist (1988). The frequency of dioecy found in a local flora therefore reflects the level of dioecy in its particular pool of families as much as, or more than, local selective factors. The positive associations of dioecy with abiotic pollination and monoecy are related to floral developmental and morphological attributes, as is the negative association with bird and bat pollination; the positive association of dioecy with climbing growth is tentatively explained in terms of differential selection for optimal resource allocation to sexual function. If rapid upward growth is at a premium in climbers and if fruit set at least temporarily inhibits growth or requires the production of thicker, more slowly growing stems to support heavy fruits, it might be advantageous to postpone femaleness. If the effect is strong, this may favor male plants.     

Dioecy,Monoecy, and Their Ecological Correlates in the Littoral Forest of Madagascar

Dioecy is a rare breeding system in flowering plants, but one that has evolved multiple times in different plant lineages. Dioecious species are commonly associated with several ecological traits, including woody habit, fleshy fruit, and small, inconspicuous flowers, although the significance of these correlations has been debated extensively. Monoecy is a breeding system that may lead to the evolution of dioecy, but ecological correlates of monoecious species have rarely been analyzed. We determined the diversity of breeding systems in the littoral forests of Madagascar and used multivariate methods to estimate which ecological traits have the strongest association with dioecy and are the best predictors of breeding systems. The Malagasy littoral forest flora is a well‐documented subset of the Malagasy flora, with 13 percent of total species diversity in an area <1 percent of the island's total. We found high levels of dioecy (18.4%) and monoecy (9.2%), similar to incidences in other tropical floras. Using multinomial logistic regression, dioecy has the strongest association with woody habit and fleshy fruit. Monoecious species have a strong association with small flowers, although this association does not hold at higher taxonomic levels. Using classification and regression tree (CART) methods, the best predictors of dioecy are woody habit and fleshy fruit; monoecy is equally predicted by fleshy and dry fruit. For the Malagasy littoral forest, both methods provide further support for the importance of woody habit and fleshy fruit in the evolution of dioecy.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

Effects of Fire on Seedling Diversity and Plant Reproduction (Sexual vs. Vegetative) in Neotropical Savannas Differing in Tree Density

Little is known about the effects of fire on the structure and species composition of Neotropical savanna seedling communities. Such effects are critical for predicting long‐term changes in plant distribution patterns in these ecosystems. We quantified richness and density of seedlings within 144 plots of 1 m² located along a topographic gradient in long‐unburned (fire protected since 1983) and recently burned (September 2005) savannas in Brazil. These savannas differ in tree density and canopy cover. Sites along the gradient, however, did not differ in species composition prior to the fire. In recently burned savannas we also evaluated the contribution of vegetative reproduction relative to sexual reproduction by quantifying richness and density of root suckers. Finally, we tested seed tolerance to pulses of high temperatures—similar to those occurring during fires on the soil surface and below—of five dominant savanna tree species. Seedlings were more abundant and diverse in unburned than in burned savannas. Seedling species composition differed among unburned and burned savannas probably reflecting early differences in root: shoot biomass allocation patterns. In recently burned savannas, root suckers were more abundant and diverse than seedlings. Relatively long exposures (>10 min) of temperatures of 90 °C reduced seed germination in all studied species suggesting a negative effect of fire on germination of seeds located at or aboveground level. Because vegetative reproduction contributes more than sexual reproduction in burned environments, frequent fires are likely to cause major shifts in species composition of Neotropical savanna plant communities, favoring clonally produced recruits along tree density/topographic gradients.     

Patch dynamics integrate mechanisms for savanna tree–grass coexistence

Many mechanisms have been suggested to explain the coexistence of woody species and grasses in savannas. However, evidence from field studies and simulation models has been mixed. Patch dynamics is a potentially unifying mechanism explaining tree–grass coexistence and the natural occurrence of shrub encroachment in arid and semi-arid savannas. A patch-dynamic savanna consists of a spatial mosaic of patches. Each patch maintains a cyclical succession between dominance of woody species and grasses, and the succession of neighbouring patches is temporally asynchronous. Evidence from empirical field studies supports the patch dynamics view of savannas. As a basis for future tests of patch dynamics in savannas, several hypotheses are presented and one is exemplarily examined: at the patch scale, realistically parameterized simulation models have generated cyclical succession between woody and grass dominance. In semi-arid savannas, cyclical successions are driven by precipitation conditions that lead to mass recruitment of shrubs in favourable years and to simultaneous collapse of shrub cohorts in drought years. The spatiotemporal pattern of precipitation events determines the scale of the savanna vegetation mosaic in space and time. In a patch-dynamic savanna, shrub encroachment is a natural, transient phase corresponding to the shrub-dominated phase during the successional cycle. Hence, the most promising management strategy for encroached areas is a large-scale rotation system of rangelands. In conclusion, patch dynamics is a possible scale-explicit mechanism for the explanation of tree–grass coexistence in savannas that integrates most of the coexistence mechanisms proposed thus far for savannas.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

Numerous predictions indicate rising CO₂ will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought‐fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire‐driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2‐million km² Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait‐based differences in fire tolerance is critical for determining the climate‐carbon‐fire feedback in tropical savanna and forest biomes.     

Multi‐scale patterns and bush encroachment in an arid savanna with a shallow soil layer

Abstract. Question: Bush encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and threatens the livelihood of many pastoralists. Can we derive a better understanding of the factors causing bush encroachment by investigating the scale dependency of patterns and processes in savannas? Location: An arid savanna in the Khomas Hochland, Namibia. Methods: Patterns of bush, grass, and soil nutrient distribution were surveyed on several scales along a rainfall gradient, with emphasis on intraspecific interactions within the dominant woody species, Acacia reficiens. Results: Savannas can be interpreted as patch‐dynamic systems where landscapes are composed of many patches (a few ha in size) in different states of transition between grassy and woody dominance. Conclusions: In arid savannas, this patchiness is driven both by rainfall that is highly variable in space and time and by inter‐tree competition. Within the paradigm of patch‐dynamic savannas, bush encroachment is part of a cyclical succession between open savanna and woody dominance. The conversion from a patch of open savanna to a bush‐encroached area is initiated by the spatial and temporal overlap of several (localized) rainfall events sufficient for Acacia germination and establishment. With time, growth and self‐thinning will transform the bush‐encroached area into a mature Acacia stand and eventually into open savanna again. Patchiness is sustained due to the local rarity (and patchiness) of rainfall sufficient for germination of woody plants as well as by plant‐soil interactions.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

Savanna woody encroachment is widespread across three continents

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta‐analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO₂, changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N‐fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.     

Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

Dioecy in the Caatinga,a Brazilian tropical dry forest: typical reproductive traits of a low frequent sexual system

Dioecy is a rare sexual system among angiosperms whose occurrence and frequency depend on the kind and geographical location of the ecosystem. We evaluate the frequency of dioecy in the Caatinga, a special semi-arid vegetation type in Brazil. We compare the floral traits and other ecological characteristics usually related to dioecy between the Caatinga and other tropical ecosystems. Dioecious species of the Caatinga were compiled based on 24 published floristic lists including most of the Caatinga domain. These lists were compared with publications comprising data on sexual systems in different kinds of vegetation in the world. Sexual system confirmation and the database of the reproductive traits of the dioecious species were made by consulting herbaria, literature, and field observations. We registered 22 dioecious species occurring in the Caatinga. The overall dioecy frequency was 2.36 %, considering all Caatinga species, and 6.91 % within woody species. Most dioecious species exhibited traits related to this sexual system (e.g., woody habit, greenish inconspicuous flowers, generalistic pollination syndrome, and zoochory). The distribution of dioecy across the Caatinga plant community exhibited a very low level of phylogenetic signal (D = ?0.145), indicating a phylogenetic structure under Brownian evolution (p = 0.671). The frequency of dioecy in the Caatinga (2.36 %) proved lower than reported for angiosperms in general (ca. 6 %) and for other tropical ecosystems. This low frequency can be related to particular ecological characteristics of the Caatinga, such as low percentage of trees >3–4 m tall (13–18.7 %), low rainfall (50 % of the Caatinga vegetation with annual rainfall lower than 750 mm), and strong seasonality (6–9 months of dry season). Dioecious species of the Caatinga exhibited typical reproductive traits for this sexual system, similar to what has been reported for dioecious species of other tropical ecosystems in the world.     

Incidence of dioecy in relation to growth form,pollination and dispersal

Summary Recent theories predict the evolution of dioecy among higher plants, in association with certain pollination and fruit dispersal traits. However, reported associations of dioecy with pollination, dispersal and growth form traits have not distinguished the effects of each trait separately, controlling for the others. Because these traits are associated among themselves, existing analyses may involve spurious or indirect correlations. This paper reports the incidence of dioecy in a subarctic and an arctic flora, and analyzes the occurrence of dioecy among vascular plants classified jointly by growth form, floral (pollination) syndrome, and fruit (dispersal) syndrome. Dioecism is no more frequent in the arctic flora as a whole, but its incidence increases northward among woody plants. This increase is associated with an increase in the proportion of woody species having small, inconspicuous flowers, and not with the syndrome of fleshy or animal dispersed fruits. Within the floras of Alaska, California, and the Northeastern US, dioecy is markedly more frequent among woody plants and among plants having small, inconspicuous flowers, and that is the only strong statistical association of dioecy for the species of these floras. When genera and families are analyzed similarly, dioecy is also associated significantly with dispersal syndrome. Thus, among angiosperms, evidence currently does not support either an uniquely strong or exclusive association of dioecy with dispersal traits, as it does for gymnosperms (Givnish 1980). It is extremely desirable to analyze the occurrence of dioecy among taxa classified jointly by all relevant ecological traits, rather than analyzing marginal distributions.     

Fire‐sensitive species dominate seed rain after fire suppression: Implications for plant community diversity and woody encroachment in the Cerrado

Woody encroachment is becoming common in tropical savannas. We studied natural seed rain and performed seed addition experiments in a Brazilian savanna that had not been burned for several decades. We found greater abundance of fire‐sensitive species in the seed rain, likely contributing to woody encroachment. Flexible fire management policies that allow for natural and prescribed fires may be required to maintain savanna diversity.     

The impact of shrub encroachment on savanna bird diversity from local to regional scale

Aim  Evidence is accumulating of a general increase in woody cover of many savanna regions of the world. Little is known about the consequences of this widespread and fundamental ecosystem structural shift on biodiversity.
Location  South Africa.
Methods  We assessed the potential response of bird species to shrub encroachment in a South African savanna by censusing bird species in five habitats along a gradient of increasing shrub cover, from grassland/open woodland to shrubland dominated by various shrub species. We also explored historical bird species population trends across southern Africa during the second half of the 20th century to determine if any quantifiable shifts had occurred that support an ongoing impact of shrub encroachment at the regional scale.
Results  At the local scale, species richness peaked at intermediate levels of shrub cover. Bird species composition showed high turnover along the gradient, suggesting that widespread shrub encroachment is likely to lead to the loss of certain species with a concomitant decline in bird species richness at the landscape scale. Finally, savanna bird species responded to changes in vegetation structure rather than vegetation species composition: bird assemblages were very similar in shrublands dominated by Acacia mellifera and those dominated by Tarchonanthus camphoratus .
Main conclusions  Shrub encroachment might have a bigger impact on bird diversity in grassland than in open woodland, regardless of the shrub species. Species recorded in our study area were associated with historical population changes at the scale of southern Africa suggesting that shrub encroachment could be one of the main drivers of bird population dynamics in southern African savannas. If current trends continue, the persistence of several southern African bird species associated with open savanna might be jeopardized regionally.     

Seed limitation of woody plants in Neotropical savannas

The failure of seeds to arrive at all suitable sites (seed limitation) greatly affects plant distribution and abundance. In contrast to tropical forests, the degree of seed limitation in Neotropical savannas is unclear because empirical studies at the community level are scarce. We estimated seed limitation of 23 woody species from annual seed rain measurements along a tree density gradient in the savannas of Central Brazil. These savannas differ in tree density and canopy cover, from closed to open savannas, and are located along shallow topographic gradients. We also studied post-dispersal seed predation and removal of 17 representative woody species, and seed viability loss over time of 12 common woody species under dry-storage conditions. Annual seed rain was lower in open (410 seeds/m²) than in closed savannas (773 seeds/m²). Average seed limitation across woody species was higher than 80% along the tree density gradient. More than 60% of seeds of the studied woody species were predated or removed within 30–45 days in all savannah types. Seeds of most common woody species (66%) lost their viability in less than 12 months of dry storage. This study shows that Neotropical savannah woody plants are strongly seed-limited because of low and poor distribution of seeds among sites, post-dispersal seed removal, and short seed longevity. The high seed limitation of tree species in Neotropical savannas, particularly in open savannas, also may contribute to maintain their relatively low tree densities and help to explain the spatial variation of tree abundance along topographic gradients.     

Patterns of plant sexual systems in subtropical evergreen broad-leaved forests in Ailao Mountains,SW China

Aims Sexual systems influence many components of the ecology and evolution of plant populations and have rarely been documented in subtropical evergreen broad-leaved forests (SEBLF). Here we report frequency distribution and ecological correlates of plant sexual systems in SEBLF at Ailao Mountains, and compare our results with that of tropical and cool temperate forests.Methods We examine the sexual systems of 703 species of woody angiosperms belonging to 103 families and 296 genera based on a comprehensive survey of SEBLF at Ailao Mountains Natural Reserve. Information of plant sexual systems and ecological traits were mainly based on published literatures and specimens as well as our field observations. The associations between plant sexual system and ecological traits were assessed with chi-square tests.Important findings Among these species, 60.2% were hermaphroditic, 15.8% were monoecious and 24.0% were dioecious. The percentage of dioecious sexual system among tree species (22.2%) in SEBLF was comparable to those of tropical tree floras, but much higher than those of temperate forests at North America. The percentage of monoecious sexual system among tree species (30.1%) in SEBLF was higher than that of tropical tree floras, but much lower than those of temperate forests at North America. Monoecy was significantly associated with the tree growth form and was relatively common in temperate genera. Dioecy was significantly associated with fleshy fruits and monoecy was significantly associated with dry fruit type in SEBLF. The high percentage of diclinous sexual systems (monoecy and dioecy) in SEBLF may be associated with the origin of the flora and the prevalence of relatively small inconspicuous flowers of the forests in the Oriental Region.     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.