Abstract: 1. Predicting and mitigating climate change effects on ecological communities is a tremendous challenge. Little attention has been given to how endemic-rich communities on isolated patches of low-nutrient soil (e.g. serpentine) will respond to climate change. 2. To address spatial factors (the isolated nature of outcrops), we incorporate habitat patchiness into species distribution models under climate change. The degree of overlap between current and future suitable habitat does not change when patchy habitats are incorporated, probably because serpentine occurs in mountainous regions where climatically and edaphically suitable regions geographically coincide. The dispersal distances required to move to newly suitable habitat are large, however, making successful migration unlikely. 3. To address how non-spatial factors affect the climate change responses of serpentine plant communities (e.g. the impacts of nutrient limitation and stress-tolerant functional traits), we conduct a literature review. Some studies suggest that serpentine communities may be at less risk than normal soil communities due to their stress-tolerant functional traits, but there is also evidence to the contrary. 4. Synthesis. Assessing climate change risk for the worlds diverse edaphic floras requires determining impacts on both special and normal soil communities. Studies are needed that use functional traits, evaluate the role of evolutionary and ecological plasticity, examine responses across spatial and temporal scales and assess the efficacy of managed relocation efforts.
Abstract: 1. Perennial plants often endure chronic loss of leaf area due to recurrent physical damage, herbivory and, for species used as non-timber forest products, due to leaf harvesting. However, little is known about functional and demographic resilience (extent and speed of recovery) of plants subjected to varying levels of chronic defoliation. 2. We used a dioecious, understorey palm (Chamaedorea elegans) to evaluate temporal trajectories and rates of recovery of leaf functional traits and vital rates (survival, growth and reproduction) after being subjected to experimental chronic defoliation regimes. 3. Pristine populations of mature C. elegans, categorized by gender (male and female), were subjected to five defoliation levels (0%, 33%, 50%, 66% or 100% of newly produced leaves) every 6 months over a period of 3 years (19972000). To evaluate recovery from defoliation, surviving palms were monitored for 3 years after the cessation of the defoliation treatment (20002003). We recorded leaf functional traits (leaf persistence, leaf production rate, leaf size and leaf area) and annual rates of mortality, growth and reproduction. 4. Cumulative effects of chronic defoliation concomitantly reduced leaf traits, survival, growth and reproduction, and this effect was stronger in female than in male palms, independent of plant size. Recovery from defoliation was faster in males than in females, but proceeded gradually overall. Survival increased first, followed by growth, while reproductive traits showed the slowest recovery rate. Recovery was independent of plant size. Notably, 3 years after defoliation treatment, the standing leaf area and probability of reproduction had not recovered to pre-defoliation levels. Additionally, we found that the occurrence of a severe drought in the first year (2000) after defoliation ceased led to decreased survival, growth and reproduction and the ability of plants to recover from defoliation. 5. Synthesis. Chronic defoliation reduces fitness components of C. elegans palms differentially between genders. Recovery is gradual and is slower and less complete in females compared with males. The lower level of resilience to chronic defoliation shown by female plants may have profound consequences for the dynamics and genetic variability of populations of tropical understorey plants undergoing prolonged defoliation. Such effects may be aggravated by severe drought episodes that are expected to increase in frequency according to global climate change predictions.
Abstract: C-4 photosynthesis is a fascinating example of parallel evolution of a complex trait involving multiple genetic, biochemical and anatomical changes. It is seen as an adaptation to deleteriously high levels of photorespiration. The current scenario for C-4 evolution inferred from grasses is that it originated subsequent to the Oligocene decline in CO2 levels, is promoted in open habitats, acts as a pre-adaptation to drought resistance, and, once gained, is not subsequently lost. We test the generality of these hypotheses using a dated phylogeny of Amaranthaceae s.l. (including Chenopodiaceae), which includes the largest number of C-4 lineages in eudicots. The oldest chenopod C-4 lineage dates back to the Eocene/Oligocene boundary, representing one of the first origins of C-4 in plants, but still corresponding with the Oligocene decline of atmospheric CO2. In contrast to grasses, the rate of transitions from C-3 to C-4 is highest in ancestrally drought resistant (salt-tolerant and succulent) lineages, implying that adaptation to dry or saline habitats promoted the evolution of C-4; and possible reversions from C-4 to C-3 are apparent. We conclude that the paradigm established in grasses must be regarded as just one aspect of a more complex system of C-4 evolution in plants in general.
Abstract: Ecologists and evolutionary biologists perceive the ecological niche as a multidimensional relationship between an organism and its environment. Yet, we know little about the degree to which multiple niche axes evolve in concert across various spatial scales to explain differences in distribution patterns and habitat specialization among lineages. Here we used contemporary phylogenetic approaches to analyze the evolution of species' distributions across multiple spatial scales in Lasthenia, a young and ecologically diverse plant clade largely occurring within the California Floristic Province, USA. Lasthenia species and subspecies range from widely distributed taxa that occupy a diversity of habitat types to locally restricted habitat endemics, including several lineages that are strongly associated with isolated ephemeral wetlands called vernal pools. We quantified the niche of Lasthenia species and subspecies at three different spatial scales: the range-wide climate niche, the habitat niche, and the within-habitat depth niche for those taxa occupying vernal pools. We incorporated phylogenetic uncertainty into our analyses by reanalyzing previously described DNA sequences in a Bayesian context and conducting all subsequent comparative analyses over the resulting posterior distribution of ultrametric phylogenetic trees. Using a biogeographic approach for ancestral habitat reconstruction, we estimated that Lasthenia lineages have undergone up to four independent transitions from strictly terrestrial habitats to a niche that incorporates semiaquatic habitats, and one of these transitions led to the subsequent proliferation of vernal pool species and subspecies. We found that the local niche axis, corresponding to the depth distribution of taxa within pools, was more phylogenetically conserved than the large-scale axes representing climatic associations. Furthermore, we did not find evidence that niche breadth estimates along different axes were consistently correlated, indicating that ecological specialization may be specific to certain niche axes rather than an overall characteristic of a species.
Abstract: Premise of the study : Pollination is a key aspect of ecosystem function in the majority of land plant communities. It is well established that many animal-pollinated plants suffer lower seed set than they are capable of, likely because of competition for pollinators. Previously, competition for pollinator services has been shown to be most intense in communities with the greatest plant diversity. In spite of the fact that community evolutionary relations have a demonstrated impact on many ecological processes, their role in competition for pollinator services has rarely been examined. Methods : In this study, we explore relations among several aspects of the surrounding plant community, including species richness, phylodiversity, evolutionary distance from a focal species, and pollen limitation in an annual insect-pollinated plant. Key results : We did not find a significant effect of species richness on competition for pollination. However, consistent with a greater role for facilitation than competition, we found that a focal species occurring in communities composed of species of close relatives, especially other members of the Asteraceae, was less pollen limited than when it occurred in communities composed of more distant relatives. Conclusions : Our results demonstrate that community phylodiversity is an important correlate of pollen limitation in this system and that it has greater explanatory power than species richness alone.
Abstract: Plant traits - the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs - determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 000 out of the world’s 300 000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation - but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models.
Abstract: Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co-occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within-community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.
Abstract: PREMISE OF THE STUDY: Pollination is a key aspect of ecosystem function in the majority of land plant communities. It is well established that many animal-pollinated plants suffer lower seed set than they are capable of, likely because of competition for pollinators. Previously, competition for pollinator services has been shown to be most intense in communities with the greatest plant diversity. In spite of the fact that community evolutionary relations have a demonstrated impact on many ecological processes, their role in competition for pollinator services has rarely been examined.
Abstract: Slender wild oat (Avena barbata) is an annual grass dominant in many grassland ecosystems in Mediterranean climate. This species has been the subject of ecological studies aimed at understanding the effect of global climate change on grassland ecosystems and the genetic basis for adaptation under varying environmental conditions. We present the sequencing and analysis of cDNA libraries constructed from leaf and root samples collected from A. barbata grown on natural soil and under varying rainfall patterns. More than 1 million expressed sequence tags (ESTs) were generated using both GS 454-FLX pyrosequencing and Sanger sequencing, and these tags were assembled into consensus sequences. We identified numerous candidate polymorphic markers in the data set, providing possibilities for linking the genomic and the existing genetic information for A. barbata. Using the digital Northern method, we showed that genes involved in photosynthesis were down-regulated under high rainfall while stress-related genes were up-regulated. We also identified a number of genes unique to the root library with unknown function. Real-time reverse transcription-PCR was used to confirm the root specificity of some of these transcripts such as two genes encoding O-methyl transferase. Also we showed differential expression of five root-specific genes under three water levels and two developmental stages. Through a combination of Sanger and 454-based sequencing technologies, we were able to generate a large set of transcribed sequences for A. barbata. This data set provides a platform for further studies of this important wild grass species.
Abstract: Aim Sustaining biological diversity requires the protection of the ecological, evolutionary and landscape-level processes that generate it. Here, we identify areas of high neoendemism in a global diversity hotspot, the California flora, using range size data and molecular-based estimates of taxon age. Location California, USA. Methods We compiled distribution and range size data for all plant taxa endemic to California and internal transcribed spacer (ITS)-based age estimates for 337 putative neoendemics (15% of the endemic flora). This information was combined to identify areas in the state with high proportions of young and restricted-range taxa. We overlaid the distribution of neoendemic hotspots on maps of currently protected lands and also explored correlations between our diversity measures and climate. Results The central coast of California, the Sierra Nevada and the San Bernardino Range contained endemics with the most restricted distributions on average, while areas in the Desert and Great Basin provinces found within the state were composed of the youngest neoendemics on average. Diversity measures that took age and range size into account shifted the estimate of highest endemic diversity in the state towards the Desert and Great Basin regions relative to simple counts of endemic species richness. Our diversity measures were poorly correlated with climate and topographic heterogeneity. Main conclusions Substantial portions of California with high levels of plant neoendemism fall outside of protected lands, indicating that additional action will be needed to preserve the geographic areas apparently associated with high rates of plant diversification. The neoendemic flora of the deserts appears particularly young in our analyses, which may reflect the relatively recent origin of desert environments within the state.
Abstract: The diversity of life is ultimately generated by evolution, and much attention has focused on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to instead remain similar over time [niche conservatism (NC)] has many consequences for the fundamental patterns and processes studied in ecology and conservation biology. Here, we describe the mounting evidence for the importance of NC to major topics in ecology (e.g. species richness, ecosystem function) and conservation (e.g. climate change, invasive species). We also review other areas where it may be important but has generally been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic interactions) and conservation (e.g. habitat modification). We summarize methods for testing for NC, and suggest that a commonly used and advocated method (involving a test for phylogenetic signal) is potentially problematic, and describe alternative approaches. We suggest that considering NC: (1) focuses attention on the within-species processes that cause traits to be conserved over time, (2) emphasizes connections between questions and research areas that are not obviously related (e.g. invasives, global warming, tropical richness), and (3) suggests new areas for research (e.g. why are some clades largely nocturnal? why do related species share diseases?).
Abstract: • Patterns of precipitation are likely to change significantly in the coming century, with important but poorly understood consequences for plant communities. Experimental and correlative studies may provide insight into expected changes, but little research has addressed the degree of concordance between these approaches. • We synthesized results from four experimental water addition studies with a correlative analysis of community changes across a large natural precipitation gradient in the United States. We investigated whether community composition, summarized with plant functional traits, responded similarly to increasing precipitation among studies and sites. • In field experiments, increased precipitation favored species with small seed size, short leaf life span and high leaf nitrogen (N) concentration. However, with increasing precipitation along the natural gradient, community composition shifted towards species with higher mean seed mass, longer leaf life span and lower leaf N concentrations. • The differences in temporal and spatial scale of experimental manipulations and natural gradients may explain these contrasting results. Our results highlight the complexity of responses to climate change, and suggest that transient dynamics may not reflect long-term shifts in functional diversity and community composition. We propose a model of community change that incorporates these differences between short- and long-term responses to climate change.
Abstract: Fire behaviour is strongly influenced by fuel load and structure; however, efforts to describe fuel patterns have largely ignored differences among species or post-fire regeneration strategies. In California chaparral, evergreen shrubs can be grouped into three post-fire regeneration strategies that correlate with a wide variety of physiological and demographic characteristics including seasonal water status and the timing of reproduction in response to fire. To test if regeneration strategy is also associated with flammability, we compared the fuel loads and structure of two post-fire seeders, Adenostoma fasciculatum and Ceanothus cuneatus, and two obligate resprouters, Heteromeles arbutifolia and Prunus ilicifolia. Species and post-fire regeneration strategies did not differ in total fuel per area, or bulk density. The proportion of fuels smaller than 6 mm in diameter differed among species, but not consistently with regeneration strategy. However, species with a post-fire seeding regeneration strategy had higher proportions of dead branches. We discuss how this difference could have arisen from evolutionary, demographic, or physiological processes.
Abstract: P>1. A number of recent studies have demonstrated that plant traits play a crucial role in determining the success or failure of species in a given environment. However, whether traits play a role in determining species’ abundance and rarity among the co-occurring species within a community remains an unresolved question. 2. To address this, we analysed the abundance of California coastal woody plant species at landscape and local scales in relation to 11 leaf, wood and seed traits. 3. At the landscape scale, we found no significant relationship between traits and abundance. In contrast, at the local scale we found significant relationships between abundance and four traits: specific leaf area (SLA), height, lumen fraction and wood density. For SLA and height, the relationship was linear; for lumen fraction, it was quadratic. For wood density, the direction of the trait-abundance relationship was dependant on the abiotic context, that is, it shifted across a gradient in soil water content. 4.Synthesis. Understanding the connections between traits and abundance is important for two reasons. First, there is an ongoing debate about the degree to which commonness and rarity are the result of drift among ecologically equivalent species or niche processes. These results suggest that there are non-random, trait-based processes affecting abundance and rarity. Secondly, species’ traits have been shown to have a strong effect on photosynthesis and decomposition rates, mediated by the abundance of the species. The connections between traits and abundance presented here are crucial for scaling from measurements of species’ traits to ecosystem-level processes.
Abstract: SUMMARY: Picante is a software package that provides a comprehensive set of tools for analyzing the phylogenetic and trait diversity of ecological communities. The package calculates phylogenetic diversity metrics, performs trait comparative analyses, manipulates phenotypic and phylogenetic data, and performs tests for phylogenetic signal in trait distributions, community structure and species interactions. AVAILABILITY: Picante is a package for the R statistical language and environment written in R and C, released under a GPL v2 open-source license, and freely available on the web (http://picante.r-forge.r-project.org) and from CRAN (http://cran.r-project.org).
Abstract: Biologists have long searched for mechanisms responsible for the increase in species richness with decreasing latitude. The strong correlation between species richness and climate is frequently interpreted as reflecting a causal link via processes linked to energy or evolutionary rates. Here, we investigate how the aggregation of clades, as dictated by phylogeny, can give rise to significant climate-richness gradients without gradients in diversification or environmental carrying capacity. The relationship between climate and species richness varies considerably between clades, regions and time periods in a global-scale phylogenetically informed analysis of all terrestrial mammal species. Many young clades show negative richness-temperature slopes (more species at cooler temperatures), with the ages of these clades coinciding with the expansion of temperate climate zones in the late Eocene. In carnivores, we find steeply positive richness-temperature slopes in clades with restricted distributions and tropical origins (e.g. cat clade), whereas widespread, temperate clades exhibit shallow, negative slopes (e.g. dog-bear clade). We show that the slope of the global climate-richness gradient in mammals is driven by aggregating Chiroptera (bats) with their Eutherian sister group. Our findings indicate that the evolutionary history should be accounted for as part of any search for causal links between environment and species richness.
Abstract: Woody stems comprise a large biological carbon fraction and determine water transport between roots and leaves; their structure and function can influence both carbon and hydrological cycles. While angiosperm wood anatomy and density determine hydraulic conductivity and mechanical strength, little is known about interrelations across many species. We compiled a global data set comprising two anatomical traits for 3005 woody angiosperms: mean vessel lumen area (Ā) and number per unit area (N). From these, we calculated vessel lumen fraction (F = ĀN) and size to number ratio (S = Ā/N), a new vessel composition index. We examined the extent to which F and S influenced potential sapwood specific stem conductivity (K(S)) and wood density (D; dry mass/fresh volume). F and S varied essentially independently across angiosperms. Variation in K(S) was driven primarily by S, and variation in D was virtually unrelated to F and S. Tissue density outside vessel lumens (D(N)) must predominantly influence D. High S should confer faster K(S) but incur greater freeze-thaw embolism risk. F should also affect K(S), and both F and D(N) should influence mechanical strength, capacitance, and construction costs. Improved theory and quantification are needed to better understand ecological costs and benefits of these three distinct dimensions.
Abstract: Environmental filtering is a fundamental process in the ecological assembly of communities. Recently developed phylogenetic tools identify patterns associated with environmental filtering across whole communities. Here we introduce a novel method that allows the detection of traits involved in the environmental filtering of species from specific clades in specific habitat types. Our approach identifies nonindependent trait/habitat/clade (THC) associations and also provides a framework for detecting clearly defined two-way trait/clade, trait/habitat, and clade/habitat associations. The THC method relies on exact binomial tests and differentiates THC associations resulting from a three-way interaction from those that are generated by one or more underlying significant two-way interactions. It can also detect THC associations for which there are no significant two-way associations (trait/habitat, trait/clade, clade/habitat). To illustrate the THC method, we examine plant pollination and dispersal traits from six habitat types in a fragmented Costa Rican landscape. Results suggest that these traits are not widely important for the environmental filtering of most clades in this landscape, but animal dispersal and insect pollination are involved in the filtering of monocots and the Piperaceae in rain forest understory.
Abstract: P> Rain forest understorey plants suffer leaf area losses due to natural causes or when leaves are harvested as non-timber forest products. The negative effects of defoliation on plant fitness can be exacerbated during periods with strong water shortage and high temperatures, typical during ENSO (El Nino Southern Oscillation) years in Mexico and Central America. At present, the isolated and combined demographic effects of ENSO events and repeated defoliation on tropical rain forest plants are poorly understood. We studied the consequences of repeated defoliation and an ENSO event on vital rates (mortality, growth, and reproduction) of the dioecious understorey palm Chamaedorea elegans. From March 1997 to March 2000 (including the 1998 ENSO year), we subjected 814 mature individuals to one of five defoliation treatments (0-100% of newly produced leaves were removed twice a year), recording mortality, growth (leaf production) and reproduction (inflorescence and seed production) every 6 months. Increasing defoliation strongly reduced reproduction but had smaller effects on growth and mortality. Among non-defoliated palms, the probability of mortality increased with light availability, likely due to drought stress during the dry season, but this was not the case for the defoliated plants, probably because leaf area removal lowered transpiration and increased the root mass-to-leaf area ratio. During the ENSO year, growth and inflorescence production were stimulated, but survivorship and seed production diminished significantly, independent of defoliation level. Synthesis. Variation in light availability and the occurrence of severe droughts can strongly affect demographic behaviour of understorey plants such as C. elegans, significantly influencing the effects of defoliation. Thus, strong episodic disturbance events (such as ENSO, insect outbreaks, strong storms, fires and landslides) should be taken into account to adequately understand the mechanisms that determine the population dynamics of forest plants and the potential for sustainable utilization of non-timber forest products.
Abstract: The concepts of niche conservatism and adaptive radiation have played central roles in the study of evolution and ecological diversification. With respect to phenotypic evolution, the two processes may be seen as opposite ends of a spectrum; however, there is no straightforward method for the comparative analysis of trait evolution that will identify these contrasting scenarios. Analysis of the rate of phenotypic evolution plays an important role in this context and merits increased attention. In this article, independent contrasts are used to estimate rates of evolution for continuous traits under a Brownian motion model of evolution. A unit for the rate of phenotypic diversification is introduced: the felsen, in honor of J. Felsenstein, is defined as an increase of one unit per million years in the variance among sister taxa of ln-transformed trait values. The use of a standardized unit of measurement facilitates comparisons among clades and traits. Rates of diversification of three functional traits (plant height, leaf size, and seed size) were estimated for four to six woody plant clades (, , , Arbutoideae, Hawaiian lobeliads, and the silversword alliance) for which calibrated phylogenies were available. For height and leaf size, rates were two to ≈300 times greater in the Hawaiian silversword alliance than in the other clades considered. These results highlight the value of direct estimates of rates of trait evolution for comparative analysis of adaptive radiation, niche conservatism, and trait diversification.
Abstract: The ranges of plants and animals are moving in response to recent changes in climate. As temperatures rise, ecosystems with 'nowhere to go', such as mountains, are considered to be more threatened. However, species survival may depend as much on keeping pace with moving climates as the climate's ultimate persistence. Here we present a new index of the velocity of temperature change (km yr(-1)), derived from spatial gradients ( degrees C km(-1)) and multimodel ensemble forecasts of rates of temperature increase ( degrees C yr(-1)) in the twenty-first century. This index represents the instantaneous local velocity along Earth's surface needed to maintain constant temperatures, and has a global mean of 0.42 km yr(-1) (A1B emission scenario). Owing to topographic effects, the velocity of temperature change is lowest in mountainous biomes such as tropical and subtropical coniferous forests (0.08 km yr(-1)), temperate coniferous forest, and montane grasslands. Velocities are highest in flooded grasslands (1.26 km yr(-1)), mangroves and deserts. High velocities suggest that the climates of only 8% of global protected areas have residence times exceeding 100 years. Small protected areas exacerbate the problem in Mediterranean-type and temperate coniferous forest biomes. Large protected areas may mitigate the problem in desert biomes. These results indicate management strategies for minimizing biodiversity loss from climate change. Montane landscapes may effectively shelter many species into the next century. Elsewhere, reduced emissions, a much expanded network of protected areas, or efforts to increase species movement may be necessary.
Abstract: This paper addresses some of the conceptual issues involved in the analysis of the age and origin of mediterranean-climate plant taxa, paying particular attention to three topics: (1) the importance of an explicit time frame in the definition of biogeographical origins, (2) the distinction between the age of traits and the age of taxa, and (3) the idea of mediterranean-type ecosystems as environmental islands. (1) In California, recent analyses demonstrate that the diversity of species derived from different biogeographical origins is significantly correlated with temperature and precipitation gradients. These patterns support the hypothesis that niche conservatism is an important factor structuring modern diversity gradients. However, depending on how far back in time one looks, a species may be assigned to different origins; future discussions of biogeographical origins need to address the appropriate time frame for analysis. (2) Past research has demonstrated distinctive trait syndromes among woody plants of the Mediterranean, Chile, California and Mexico, and proposed that the syndromes are associated with lineages of different age in these floras. Reanalysis of individual traits demonstrates greater variability among regions than previously reported. The classification of plants into ‘old’ and ‘new’ genera is re-evaluated, and it is suggested that greater attention be paid to the age of traits, rather than to the age of taxa, especially at an arbitrary rank such as genus. (3) The idea of mediterranean-climate regions as ‘climatic islands’ is examined. Space-time diagrams of climate enable one to view the emergence of distinctive climatic regions in a continental context. The terms ‘synclimatic’ and ‘anticlimatic’ are proposed, referring to migration routes that parallel climate contours in space and time versus those that cross contours (including the case of geographic stasis in the face of climate change), respectively. Mediterranean-climate regions have served as important case studies in plant ecology and evolution, and merit continued close examination in the light of continued advances in phylogenetics and palaeoecology.
Abstract: Environmental filtering is a fundamental process in the ecological assembly of communities. Recently developed phylogenetic tools identify patterns associated with environmental filtering across whole communities. Here we introduce a novel method that allows the detection of traits involved in the environmental filtering of species from specific clades in specific habitat types. Our approach identifies nonindependent trait/habitat/clade (THC) associations and also provides a framework for detecting clearly defined two-way trait/clade, trait/habitat, and clade/habitat associations. The THC method relies on exact binomial tests and differentiates THC associations resulting from a three-way interaction from those that are generated by one or more underlying significant two-way interactions. It can also detect THC associations for which there are no significant two-way associations (trait/habitat, trait/clade, clade/habitat). To illustrate the THC method, we examine plant pollination and dispersal traits from six habitat types in a fragmented Costa Rican landscape. Results suggest that these traits are not widely important for the environmental filtering of most clades in this landscape, but animal dispersal and insect pollination are involved in the filtering of monocots and the Piperaceae in rain forest understory.
Abstract: The flora of California, a global biodiversity hotspot, includes 2387 endemic plant taxa. With anticipated climate change, we project that up to 66% will experience >80% reductions in range size within a century. These results are comparable with other studies of fewer species or just samples of a region's endemics. Projected reductions depend on the magnitude of future emissions and on the ability of species to disperse from their current locations. California's varied terrain could cause species to move in very different directions, breaking up present-day floras. However, our projections also identify regions where species undergoing severe range reductions may persist. Protecting these potential future refugia and facilitating species dispersal will be essential to maintain biodiversity in the face of climate change.
Abstract: Leaf mass per area (LMA) is one of the most widely measured of all plant functional traits. In deciduous forests, there is similarity between plastic and evolutionary responses of LMA to light gradients. In evergreens, however, LMA is lower in shaded than sunlit individuals of the same species, whereas shade-tolerant evergreens have higher LMA than light-demanders grown under the same conditions. We suggest that this pattern of 'counter-gradient variation' results from some combination of (i) close evolutionary coordination of LMA with leaf lifespan, (ii) selection for different leaf constitutions (relative investment in cell walls versus cell contents) in sun and shade environments and/or (iii) constraints on plasticity as a result of genetic correlations between phenotypes expressed in sun and shade.
Abstract: It is debated whether species-level differences in ecological strategy, which play a key role in much of coexistence theory, are important in structuring highly diverse communities. We examined the co-occurrence patterns of over 1100 tree species in a 25-hectare Amazonian forest plot in relation to field-measured functional traits. Using a null model approach, we show that co-occurring trees are often less ecologically similar than a niche-free (neutral) model predicts. Furthermore, we find evidence for processes that simultaneously drive convergence and divergence in key aspects of plant strategy, suggesting that at least two distinct niche-based processes are occurring. Our results show that strategy differentiation among species contributes to the maintenance of diversity in one of the most diverse tropical forests in the world.
Abstract: Most studies of plant community assembly have focused on how the abiotic aspects of a habitat (e.g. soil moisture or mineral composition) or direct interactions among plants in a community (e.g. competition for space or nutrients) influence which species establish and persist, but they have tended to neglect indirect interactions such as those mediated by pollinators. We address three types of plant-pollinator interactions--filtering, facilitation and competitive exclusion--and their predicted impacts on communities. The few available studies that address how pollinator-mediated interactions limit or promote plant species establishment and persistence provide support for many of these predictions. An integrated framework for understanding plant community assembly needs to incorporate abiotic and biotic interactions, including plant-pollinator and other plant-animal interactions.
Abstract: A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.
Abstract: MOTIVATION: The increasing availability of phylogenetic and trait data for communities of co-occurring species has created a need for software that integrates ecological and evolutionary analyses. Capabilities: Phylocom calculates numerous metrics of phylogenetic community structure and trait similarity within communities. Hypothesis testing is implemented using several null models. Within the same framework, it measures phylogenetic signal and correlated evolution for species traits. A range of utility functions allow community and phylogenetic data manipulation, tree and trait generation, and integration into scientific workflows. Availability: Open source at: http://phylodiversity.net/phylocom/.
Abstract: Taxa co-occurring in communities often represent a nonrandom sample, in phenotypic or phylogenetic terms, of the regional species pool. While heuristic arguments have identified processes that create community phylogenetic patterns, further progress hinges on a more comprehensive understanding of the interactions between underlying ecological and evolutionary processes. We created a simulation framework to model trait evolution, assemble communities (via competition, habitat filtering, or neutral assembly), and test the phylogenetic pattern of the resulting communities. We found that phylogenetic community structure is greatest when traits are highly conserved and when multiple traits influence species membership in communities. Habitat filtering produces stronger phylogenetic structure when taxa with derived (as opposed to ancestral) traits are favored in the community. Nearest-relative tests have greater power to detect patterns due to competition, while total community relatedness tests perform better with habitat filtering. The size of the local community relative to the regional pool strongly influences statistical power; in general, power increases with larger pool sizes for communities created by filtering but decreases for communities created by competition. Our results deepen our understanding of processes that contribute to phylogenetic community structure and provide guidance for the design and interpretation of empirical research.
Abstract: Plant functional traits vary both along environmental gradients and among species occupying similar conditions, creating a challenge for the synthesis of functional and community ecology. We present a trait-based approach that provides an additive decomposition of species' trait values into alpha and beta components: beta values refer to a species' position along a gradient defined by community-level mean trait values; alpha values are the difference between a species' trait values and the mean of co-occurring taxa. In woody plant communities of coastal California, beta trait values for specific leaf area, leaf size, wood density and maximum height all covary strongly, reflecting species distributions across a gradient of soil moisture availability. Alpha values, on the other hand, are generally not significantly correlated, suggesting several independent axes of differentiation within communities. This trait-based framework provides a novel approach to integrate functional ecology and gradient analysis with community ecology and coexistence theory.
Abstract: Chaparral shrubs in California experience cool, wet winters and hot, dry summers characteristic of mediterranean-type climates; by contrast, morphologically similar close relatives in central Mexico experience summer rainfall. A comparison of closely related species pairs was conducted to examine whether evolutionary divergences in plant hydraulic conductivity were associated with contrasting seasonality of precipitation. Six species pairs in Santa Barbara, California and Tehuacan, Mexico were chosen to test for repeated directional divergences across the habitat contrast. Additionally, evolutionary correlations were examined using phylogenetically independent contrasts (PICs) among a suite of hydraulic traits, including stem- and leaf-specific conductivity, resistance to embolism, wood density, inverse Huber value, and minimum seasonal water potential. Leaf-specific conductivity was generally higher in California, but for most hydraulic traits the species pairs exhibited varied evolutionary trajectories across the climate contrast. A significant correlation was found between divergences in xylem resistance to embolism and minimum seasonal water potential, but no evolutionary trade-off was found between resistance and stem conductivity. Higher leaf-specific conductivity may be adaptive in California, where soil and atmospheric droughts coincide during summer months. This response is consistent with a hydraulic strategy of high leaf water supply under high evaporative demand to prevent excessive drops in water potential.
Abstract: Aim To provide the first global quantification of the slope and shape of the latitudinal gradient in seed mass, and to determine whether global patterns in seed mass are best explained by growth form, vegetation type, seed dispersal syndrome, or net primary productivity (NPP). Location Global. Methods We collected seed mass data for 11,481 species x site combinations from around the world. We used regression to describe the latitudinal gradient in seed mass, then applied general linear models to quantify the relative explanatory power of each of the variables hypothesized to underlie the latitudinal gradient in seed size. Results There is a 320-fold decline in geometric mean seed mass between the equator and 60 degrees. This decline is not linear. At the edge of the tropics, there is a sudden 7-fold drop in mean seed mass. The strongest correlates of the latitudinal gradient in seed mass are plant growth form, and vegetation type, followed by dispersal syndrome and NPP. A model including growth form, vegetation type, dispersal syndrome and NPP explains 51% of the variation in seed mass. Latitude explains just 0.2% of the residual variation from this model. Main Conclusions This is the first demonstration of a major decrease in seed size at the edge of the tropics. This drop in seed mass is most closely correlated with changes in plant growth form and vegetation type. This suggests that the drop in seed mass might be part of a sudden change in plant strategy at the edge of the tropics.
Abstract: We propose research to fill key gaps in the areas of population and community ecology, based on a National Science Foundation workshop identifying funding priorities for the next 5-10 years. Our vision for the near future of ecology focuses on three core areas: predicting the strength and context-dependence of species interactions across multiple scales; identifying the importance of feedbacks from individual interactions to ecosystem dynamics; and linking pattern with process to understand species coexistence. We outline a combination of theory development and explicit, realistic tests of hypotheses needed to advance population and community ecology.
Abstract: 1. Tropical rain forest trees have been classified according to the importance of canopy disturbances for their life histories. To assess gap-dependence vs. shade tolerance, we propose using demographic analysis of the degree to which gaps are critical for population growth.2. Population growth is the composite of the successes and failures of individuals experiencing different life-cycle pathways (i.e. gap, building, and mature forest patches). To assess the contribution of each pathway, it is necessary to study patch dynamics.3. We use stage and patch structured matrix models and loop analysis to assess the life-history strategies of two Neotropical mangrove trees, Avicennia germinans and Rhizophora mangle. We use data from field studies in Lake Maracaibo, Venezuela, that were specifically designed to parameterize these models.4. In R. mangle, seedling to juvenile progression was limited to high light patches (> 20% canopy openness), even though they comprised < 10% of the area, and population growth depended entirely on gaps. A. germinans seedlings progressed to juveniles from all light patches, and progression rates increased with light level. However, low and medium light patches (< 20% openness) were most important to population growth because they were most frequent. We suggest that R. mangle is gap-dependent while A. germinans is shade-promoted (i.e. shaded patches are demographically more important to A. germinans population growth even though seedling progression rates increase with canopy openness).5. Previous studies considered R. mangle shade-tolerant because its propagules can establish and initially survive in the shade, but our classification was based on its entire life cycle. For A. germinans, the important measure was not the growth response to light, but the frequency and demographic importance of different light patches.6. Synthesis: These examples show that observations of a plant’s germination biology or photosynthetic/growth response to light do not necessarily translate into demographic behaviour because physiology can be decoupled from demography by patch dynamics. Thus, it is critical to consider the entire life cycle in the context of forest patch dynamics when assessing the life-history strategies of tropical trees.
Abstract: When ecologically important plant traits are correlated they may be said to constitute an ecological 'strategy' dimension. Through identifying these dimensions and understanding their inter-relationships we gain insight into why particular trait combinations are favoured over others and into the implications of trait differences among species. Here we investigated relationships among several traits, and thus the strategy dimensions they represented, across 2134 woody species from seven Neotropical forests.
Abstract: Eukaryotic genome size varies over five orders of magnitude; however, the distribution is strongly skewed toward small values. Genome size is highly correlated to a number of phenotypic traits, suggesting that the relative lack of large genomes in eukaryotes is due to selective removal. Using phylogenetic contrasts, we show that the rate of genome size evolution is proportional to genome size, with the fastest rates occurring in the largest genomes. This trend is evident across the 20 major eukaryotic clades analyzed, indicating that over long time scales, proportional change is the dominant and universal mode of genome-size evolution in eukaryotes. Our results reveal that the evolution of eukaryotic genome size can be described by a simple proportional model of evolution. This model explains the skewed distribution of eukaryotic genome sizes without invoking strong selection against large genomes.
Abstract: We have studied the interactive effects of salinity and light on Avicennia germinans mangrove seedlings in greenhouse and field experiments. We hypothesized that net photosynthesis, growth, and survivorship rates should increase more with an increase in light availability for plants growing at low salinity than for those growing at high salinity. This hypothesis was supported by our results for net photosynthesis and growth. Net daily photosynthesis did increase more with increasing light for low-salinity plants than for high-salinity plants. Stomatal conductance, leaf-level transpiration, and internal CO(2) concentrations were lower at high than at low salinity. At high light, the ratio of leaf respiration to assimilation was 2.5 times greater at high than at low salinity. Stomatal limitations and increased respiratory costs may explain why, at high salinity, seedlings did not respond to increased light availability with increased net photosynthesis. Seedling mass and growth rates increased more with increasing light availability at low than at high salinity. Ratios of root mass to leaf mass were higher at high salinity, suggesting that either water or nutrient limitations may have limited seedling growth at high salinity in response to increasing light. The interactive effects of salinity and light on seedling size and growth rates observed in the greenhouse were robust in the field, despite the presence of other factors in the field--such as inundation, nutrient gradients, and herbivory. In the field, seedling survivorship was higher at low than at high salinity and increased with light availability. Interestingly, the positive effect of light on seedling survivorship was stronger at high salinity, indicating that growth and survivorship rates are decoupled. In general, this study demonstrates that environmental effects at the leaf-level also influence whole plant growth in mangroves.
Abstract: In the course of an adaptive radiation, the evolution of niche parameters is of particular interest for understanding modes of speciation and the consequences for coexistence of related species within communities. We pose a general question: In the course of an evolutionary radiation, do traits related to within-community niche differences (alpha niche) evolve before or after differentiation of macrohabitat affinity or climatic tolerances (beta niche)? Here we introduce a new test to address this question, based on a modification of the method of independent contrasts. The divergence order test (DOT) is based on the average age of the nodes on a tree, weighted by the absolute magnitude of the contrast at each node for a particular trait. The comparison of these weighted averages reveals whether large divergences for one trait have occurred earlier or later in the course of diversification, relative to a second trait; significance is determined by bootstrapping from maximum-likelihood ancestral state reconstructions. The method is applied to the evolution of Ceanothus, a woody plant group in California, in which co-occurring species exhibit significant differences in a key leaf trait (specific leaf area) associated with contrasting physiological and life history strategies. Co-occurring species differ more for this trait than expected under a null model of community assembly. This alpha niche difference evolved early in the divergence of two major subclades within Ceanothus, whereas climatic distributions (beta niche traits) diversified later within each of the subclades. However, rapid evolution of climate parameters makes inferences of early divergence events highly uncertain, and differentiation of the beta niche might have taken place throughout the evolution of the group, without leaving a clear phylogenetic signal. Similar patterns observed in several plant and animal groups suggest that early divergence of alpha niche traits might be a common feature of niche evolution in many adaptive radiations.
Abstract: We hypothesized that salinity and light interactively affect mangroves, such that net photosynthesis, growth, and survivorship rates increase more with increase in light availability at low than high salinity. Using greenhouse and field experiments, we determined that net photosynthesis, growth rates, and size increased more with light at low than high salinity. At high salinity, the ratio of leaf respiration to assimilation increased fourfold, suggesting that salinity may have contributed to declines in net photosynthesis. Stomatal conductance, leaf-level transpiration, and internal CO2 concentrations were lower at high salinity. Ratios of root mass to leaf mass were higher at high salinity. Stomatal limitations and increased respiratory costs may explain why at high salinity, the seedlings did not respond to increased light availability with increased net photosynthesis. Increased root mass relative to leaf mass suggests that at high salinity, either water or nutrient limitations may have prevented the seedlings from increasing growth with increasing light availability. At both low- and high-salinity zones in the field, seedling survivorship increased with light availability, and the effect of light was stronger at low salinity. However, at low light, survivorship was higher at high than low salinity, indicating that there may be a trade-off between survivorship and growth. The interactive effects observed in the greenhouse were robust in the field, despite the presence of other factors in the field such as inundation and nutrient gradients and herbivory. This study provides a robust test of the hypothesis that salinity and light interactively effect mangrove seedling performance.
Abstract: dThe minimum seasonal water potential a plant experiences, psi(min), provides an important measure of plant water status, as it reflects the maximum water deficit that leaves and xylem must tolerate to maintain physiological activity. psi(min) also acts as a selective force on xylem structure which, in turn, generates correlations between psi(min) and numerous hydraulic traits. This review focuses on the ecological relevance Of psi(min) as a reflection of overall plant hydraulic strategy. The focus is on plant functional strategies with respect to soil drought, but we conclude with preliminary findings on the role of atmospheric drought.
Abstract: Community assembly theory suggests that two processes affect the distribution of trait values within communities: competition and habitat filtering. Within a local community, competition leads to ecological differentiation of coexisting species, while habitat filtering reduces the spread of trait values, reflecting shared ecological tolerances. Many statistical tests for the effects of competition exist in the literature, but measures of habitat filtering are less well-developed. Here, we present convex hull volume, a construct from computational geometry, which provides an n-dimensional measure of the volume of trait space occupied by species in a community. Combined with ecological null models, this measure offers a useful test for habitat filtering. We use convex hull volume and a null model to analyze California woody-plant trait and community data. Our results show that observed plant communities occupy less trait space than expected from random assembly, a result consistent with habitat filtering.
Abstract: Human-altered landscapes dominate the planet, yet little is known about their capacity to sustain plant functional diversity. Most conservation-orientated studies of such landscapes focus on species diversity, whereas less attention is given to functional traits and their conservation.We examine the functional diversity of herbaceous and shrubby plant communities in three forest habitats (understorey, tree-fall gaps and riverbanks) and three deforested habitats (pasture, roadside vegetation and pasture riverbanks), each replicated in three human-dominated landscapes in southern Costa Rica. We focus on six categorical traits related to forest regeneration, reproduction and dispersal: pollination mechanism, dispersal mechanism, growth form, fruit type, fruit size and seed size.We compared trait state richness and composition of each trait in forested and deforested habitats and how three pollination states (bat, bird and bee pollination) and three dispersal states (fur, bird and monkey dispersal) of conservation interest were distributed across these landscapes.Only one trait state was missing from forest, and none was missing from deforested habitats. Understorey and pasture were consistently trait state poor. Forested and deforested plots differed in trait state composition for all traits. Pasture riverbanks and road verges were compositionally similar to forest riverbanks and tree-fall gaps, for multiple traits. There were more compositional similarities between forested and deforested habitat types when abundance of individuals with a trait state was used as the basis for similarity measures than when the number of species with each trait state was used. Bat-, bird- and bee-pollinated plants and plants with bird- and monkey-dispersed fruits were most common in forest and pasture riverbanks whereas species with fur-dispersed seeds were more common in all deforested habitats.Functional diversity patterns were inconsistent across habitat types and locations but overall functional similarity was high between forested and deforested communities. Patterns of functional diversity were far more variable between habitats and landscapes than the consistent patterns found previously for species diversity.
Abstract: Despite growing attention to how human activities alter plant communities, little is known about the ecosystem consequences of these changes. We explore the relationship between species and functional diversity of herbaceous and shrubby plant communities in forested and deforested habitats in three Neotropical landscapes. We focus on six traits: pollination mechanism, dispersal mechanism, growth form, fruit type, fruit size, and seed size. We ask: (1) What is the relationship between species richness and functional diversity (trait state richness)? (2) Do species/functional diversity relationships differ between forested and deforested habitats? and (3) Are observed species/functional diversity patterns more consistent with ecological filtering or differentiation-based assembly processes? We show that species richness is often a weak surrogate for functional diversity, depending on the trait. Species/functional diversity relationships differ significantly between forested and deforested habitats, but the nature of-differences is trait dependent. Dispersal mechanism and fruit type number increased more rapidly in deforested than forested habitats, but the opposite was true for most other traits. Using a null model, we found evidence of ecological filtering for most traits in both habitats. Results demonstrate that deforested habitats do not necessarily contain lower functional diversity than forest but that the ecological assembly processes influencing community function in deforested communities differ dramatically from forest.
Abstract: Recent developments in community models emphasize the importance of incorporating stochastic processes (e.g. ecological drift) in models of niche-structured community assembly. We constructed a finite, spatially explicit, lottery model to simulate the distribution of species in a one-dimensional landscape with an underlying gradient in environmental conditions. Our framework combines the potential for ecological drift with environmentally-mediated competition for space in a heterogeneous environment. We examined the influence of niche breadth, dispersal distances, community size (total number of individuals) and the breadth of the environmental gradient on levels of species and functional trait diversity (i.e. differences in niche optima). Three novel results emerge from this model: (1) niche differences between adjacent species (e.g. limiting similarity) increase in smaller communities, because of the interaction of competitive effects and finite population sizes; (2) immigration from a regional species pool, stochasticity and niche-assembly generate a bimodal distribution of species residence times (’transient’ and ’resident’) under a heterogeneous environment; and (3) the magnitude of environmental heterogeneity has a U-shaped effect on diversity, because of shifts in species richness of resident vs. transient species. These predictions illustrate the potential importance of stochastic (although not necessarily neutral) processes in community assembly.
Abstract: Many woody plant species that depend upon fire-cued seed germination lack the ability to resprout. As the ability to resprout is widely assumed to be the ancestral condition in most plant groups, the failure to sprout is an evolutionary derived trait. Models for the evolutionary loss of sprouting assume a trade-off between seedling success and vegetative resprouting ability of adults. Such models require higher seedling success rates in nonsprouters than in sprouters. On the other hand, there seem to be few a priori reasons why a strong sprouter might not also have highly competitive post-fire seedlings. To test the hypothesis that nonsprouting plants have higher growth rates and/or drought survival, we grew seedlings of Ceanothus tomentosus from sprouting and nonsprouting populations in a common garden experiment. Each of these C. tomentosus populations was paired with a sympatric Ceanothus species that differed in resprouting ability. Sprouters exhibited greater allocation to root carbohydrate storage than did nonsprouters, but overall relative growth rates did not differ. Nonsprouters had earlier onset of flowering. These results provide mixed support for models of a sprouting/nonsprouting allocation trade-off.
Abstract: Improved phylogenies and the accumulation of broad comparative data sets have opened the way for phylogenetic analyses to trace trait evolution in major groups of organisms. We arrayed seed mass data for 12,987 species on the seed plant phylogeny and show the history of seed size from the emergence of the angiosperms through to the present day. The largest single contributor to the present-day spread of seed mass was the divergence between angiosperms and gymnosperms, whereas the widest divergence was between Celastraceae and Parnassiaceae. Wide divergences in seed size were more often associated with divergences in growth form than with divergences in dispersal syndrome or latitude. Cross-species studies and evolutionary theory are consistent with this evidence that growth form and seed size evolve in a coordinated manner.
Abstract: We used correlated divergence analysis to determine which factors have been most closely associated with changes in seed mass during seed plant evolution. We found that divergences in seed mass have been more consistently associated with divergences in growth form than with divergences in any other variable. This finding is consistent with the strong relationship between seed mass and growth form across present-day species and with the available data from the paleobotanical literature. Divergences in seed mass have also been associated with divergences in latitude, net primary productivity, temperature, precipitation, and leaf area index. However, these environmental variables had much less explanatory power than did plant traits such as seed dispersal syndrome and plant growth form.
Abstract: In this computational study, we examined optimal reproductive allocation schedules in annual plants whose season lengths vary in predictability. We discuss relationships among season-length predictability, the form of the optimal allocation schedule, the degree of plasticity reflected in the optimal reaction norm, and the competitive consequences of plasticity and bet-hedging. We used an evolutionary algorithm to search the allocation-schedule space for optima, given different distributions of season length. The resulting schedules maximize geometric-mean fecundity under their selecting distributions. We then examined the relative fitness of these schedules in simulated competition among reaction norms optimized for different degrees of season-length predictability. Gradedness of optimal schedules decreases with increasing season-length predictability, and reaction norms comprising highly graded schedules reflect lesser plasticity than norms comprising schedules that are less graded. In simulations, competitively successful genotypes were those that reflected plasticity appropriate to the season-length predictability. Informational constraints in the form of low season-length predictability select for low plasticity and high bet-hedging in allocation. Because an environmental cue must mediate the relationship between environment and fitness, plasticity in reproductive allocation ought to be understood not as a direct response to the selective environment, but rather to cues that are correlated with relevant environmental parameters.
Abstract: Small leaves and low specific leaf area (SLA) have long been viewed as adaptations to Mediterranean-type climates in many species of evergreen woody plants. However, paleobotanical and floristic evidence suggests that in many cases these traits originated prior to the advent of the summer-drought climate regime. In this study, molecular phylogenies and ancestral state reconstructions were used to test the hypothesis of adaptive leaf evolution in 12 lineages of evergreen shrubs in the California chaparral. Across all lineages there was a small but significant shift toward lower SLA, but there were no trends in leaf size evolution. For individual lineages, adaptive changes were detected in only three cases for SLA and in one case for leaf size. Three of these cases of evolutionary change were observed in taxa derived from cool temperate ancestors (e.g., Heteromeles). In contrast, most lineages originating from subtropical ancestors exhibited relative stasis in leaf trait evolution (e.g., Ceanothus). The absence of change suggests that ancestors of chaparral taxa had already acquired appropriate traits that contributed to their success under Mediterranean-type climates. These results illustrate how biogeographic history may influence patterns of trait evolution and adaptation and highlight the contribution of ecological sorting processes to the assembly and functional ecology of regional biotas.
Abstract: Evolutionary diversification of a phenotypic trait reflects the tempo and mode of trait evolution, as well as the phylogenetic topology and branch lengths. Comparisons of trait variance between sister groups provide a powerful approach to test for differences in rates of diversification, controlling for differences in clade age. We used simulation analyses under constant rate Brownian motion to develop phylogenetically based F-tests of the ratio of trait variances between sister groups. Random phylogenies were used for a generalized evolutionary null model, so that detailed internal phylogenies are not required, and both gradual and speciational models of evolution were considered. In general, phylogenetically structured tests were more conservative than corresponding parametric statistics (i.e., larger variance ratios are required to achieve significance). The only exception was for comparisons under a speciational evolutionary model when the group with higher variance has very low sample size ( number of species). The methods were applied to a large data set on seed size for 1976 species of California flowering plants. Seven of 37 sister-group comparisons were significant for the phylogenetically structured tests ( compared to 12 of 37 for the parametric F-test). Groups with higher diversification of seed size generally had a greater diversity of fruit types, life form, or life history as well. The F-test for trait variances provides a simple, phylogenetically structured approach to test for differences in rates of phenotypic diversification and could also provide a valuable tool in the study of adaptive radiations.
Abstract: The study of interspecific variation in plant ecological strategies has revealed suites of traits associated with leaf life span and with maximum levels of water deficit (measured as leaf water potentials). Here, the relationship between these sets of traits was examined in a study of 20 co-occurring chaparral shrubs that vary in leaf habit, rooting depth, and regeneration strategies. Leaf life span (LLS) and minimum seasonal water potentials (psi(min)) were not significantly correlated, suggesting that they are associated with independent aspects of functional variation. Multiple regression analyses of a large suite of physiological, functional, and phenological attributes in relation to these two "anchor traits" supported this view. Short LLS was significantly associated with high specific leaf area, hi-h carbon assimilation and leaf nitrogen (per mass), early onset of growth, and a multistemmed, short stature growth form. This suite of traits was also associated with opportunistic regeneration following physical disturbance. Area-based gas exchange was not tightly linked to LLS. Low psi(min) (i.e., greater water deficit) was associated with high wood density, small vessel diameters, thin twigs, low leaf area: sapwood area ratios, and early onset of leaf abscission. Among the evergreen species, this suite of traits was most characteristic of post-fire seeders, which depend on high drought tolerance for post-fire regeneration of seedlings. Plant stature was the only trait associated with both the LLS axis and the psi(min) axis of functional variation. A two-dimensional strategy space, approximately defined by LLS and psi(min), can be used to distinguish contrasting strategies of drought tolerance vs. avoidance, and alternative modes of regeneration following fire and other disturbance. This conceptual scheme illustrates the strength of a trait-based approach to defining plant strategies in relation to resource availability and disturbance.
Abstract: Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.
Abstract: Closely related species that occur together in communities and experience similar environmental conditions are likely to share phenotypic traits because of the process of environmental filtering. At the same time, species that are too similar are unlikely to co-occur because of competitive exclusion. In an effort to explain the coexistence of 17 oak species within forest communities in North Central Florida, we examined correlations between the phylogenetic relatedness of oak species, their degree of co-occurrence within communities and niche overlap across environmental gradients, and their similarity in ecophysiological and life-history traits. We show that the oaks are phylogenetically overdispersed because co-occurring species are more distantly related than expected by chance, and oaks within the same clade show less niche overlap than expected. Hence, communities are more likely to include members of both the red oak and the white + live oak clades than only members of one clade. This pattern of phylogenetic overdispersion arises because traits important for habitat specialization show evolutionary convergence. We hypothesize further that certain conserved traits permit coexistence of distantly related congeners. These results provide an explanation for how oak diversity is maintained at the community level in North Central Florida.
Abstract: We used pairs of congeneric shrub species from contrasting habitats to test for repeated evolutionary divergence in leaf-stem allometry and shoot hydraulic architecture in response to water availability. Allometric relationships and mean ratios between leaf size (individual and total area and mass per shoot) and stem cross-sectional area were compared between habitats using six species pairs representing three genera (Arctostaphylos, Baccharis, Ceanothus). We measured correlations among evolutionary changes in allometric, morphological, and physiological traits using phylogenetic independent contrasts. Allometric analysis revealed habitat differences: slopes were homogeneous among species (=1.46), but the more mesic-adapted species generally supported more leaf area at a common stem cross-sectional area. Reducing bivariate allometry to a ratio obscured this pattern because ratios varied with stem size, which was unrelated to habitat. Mean individual leaf size also was not correlated with either water availability or leaf-stem allometry. Stem hydraulic conductivity was generally lower in the xeric-adapted species of each pair, and its evolution mirrored changes in shoot allometry. This study provides evidence for repeated evolutionary divergence in shoot allometry and hydraulic architecture associated with water availability and demonstrates the importance of shoot allometry to water relations, independent of leaf size.
Abstract: We examined whether increased high temperature photosynthetic thermal tolerance (PT), reduced specific leaf area (SLA) and reduced leaf size represent correlated and convergent adaptations for recently diverged Encelia, Salvia, Atriplex and Eriogonum congeneric species pairs from contrasting thermal and water environments (the Mojave Desert and coastal California). We also studied whether variation in PT is associated with inducible small heat shock protein expression (sHsp).Traits were measured in a common environment (CE) and in the field to partition effects of phenotypic plasticity and genetic divergence.We found little evidence for convergent adaptation of PT (CE measurements). Field measurements revealed significant plasticity for PT, which was also associated with increased sHsp expression. Compared to coastal congeners desert species had lower SLA in the CE. These differences were magnified in the field. There was a negative correlation between SLA and PT. Desert species also tended to have smaller leaves both in the CE and in the field.SLA and leaf size reductions represent repeated evolutionary divergences and are perhaps convergent adaptations for species radiating into the desert, while PT is highly plastic and shows little evidence for convergent adaptation in the congeneric species pairs we studied.
Abstract: We used pairs of congeneric shrub species from contrasting habitats to test for repeated evolutionary divergence in leaf-stem allometry and shoot hydraulic architecture in response to water availability. Allometric relationships and mean ratios between leaf size (individual and total area and mass per shoot) and stein cross-sectional area were compared between habitats using six species pairs representing three genera (Arctostaphylos, Baccharis, Ceanothus). We measured correlations among evolutionary changes in allometric. morphological, and physiological traits using phylogenetic independent contrasts. Allometric analysis revealed habitat differences: slopes were homogeneous among species (= 1.46), but the more mesic-adapted species generally supported more leaf area at a common stem cross-sectional area. Reducing bivariate allometry to a ratio obscured this pattern because ratios varied with stem size, which was unrelated to habitat. Mean individual leaf size also was not correlated with either water availability or leaf stem allometry. Stem hydraulic conductivity was generally lower in the xeric-adapted species of each pair, and its evolution mirrored changes in shoot allometry. This study provides evidence for repeated evolutionary divergence in shoot allometry and hydraulic architecture associated with water availability and demonstrates the importance of shoot allometry to water relations, independent of leaf size.
Abstract: The widespread correspondence between phenotypic variation and environmental conditions, the “fit†of organisms to their environment, reflects the adaptive value of plant functional traits. Several processes contribute to these patterns: plasticity, ecological sorting, and adaptive evolution. This article addresses the importance of ecological sorting processes (community assembly, migration, habitat tracking, etc.) as primary causes of functional trait distributions at the local and landscape level. In relatively saturated communities, plants will establish and regenerate in environments to which they are well adapted, so their distributions, and the distributions of associated functional traits, will reflect the distribution of optimal or near-optimal environmental conditions in space and time. The predicted evolutionary corollary of this process is that traits related to habitat occupancy, e. g., environmental tolerances, will be under stabilizing selection. This process contributes to the widely observed pattern of phylogenetic niche conservatism, i. e., ecological and phenotypic similarities of closely related species. Evidence for niche conservatism in plants is reviewed. Based on Jackson and Overpeck’s concept of the realized environment, I propose three scenarios in which a species’ distributional responses to environmental conditions will lead to a “mismatch†between its environmental tolerances and the environments it occupies, thus creating opportunities for adaptive evolution: (1) the colonization of “environmental islands†(habitats that are discontinuous in niche space) that require large adaptive shifts in tolerance of one or more environmental factors; (2) the persistence of “trailing-edge†populations in species tracking changing climate, if barriers to dispersal of competitors prevent competitive exclusion in the deteriorating conditions; and (3) responses to changes in the realized environment in multidimensional niche space, in which species are predicted to track environmental factors for which they exhibit narrow tolerances and exhibit adaptive evolutionary response along axes where they exhibit greater niche breadth. These three scenarios provide a conceptual framework that emphasizes the role of ecological sorting processes and stabilizing selection as the context for understanding opportunities for adaptive evolution in heterogeneous and changing environments.
Abstract: We analyzed to what extent and by what mechanisms plants of the tropical understory palm Chamaedorea elegans are able to mitigate the negative effects of defoliation on performance (i.e., plant size, total growth, leaf lamina growth, and reproduction) and how this is related to light availability. For this purpose we developed a new approach that allowed us to quantify the performance of defoliated plants relative not only to the performance of undamaged plants, but also relative to the estimated performance of hypothetical defoliated plants that do not exhibit any. mechanisms of compensatory growth. The latter provides a way to quantify the adaptive value of compensation with reference to a hypothetical noncompensating alternative state. C. elegans plants were grown in a greenhouse at two light levels (5% and 16% of natural daylight) and subjected to five defoliation treatments (a control and four levels of defoliation). Defoliation was repeated every three months. Growth analysis revealed that defoliated plants allocated considerably more mass to the production of leaf laminas (f(lam)) than control plants, at the expense of allocation to other organs, particularly reproductive structures. Average growth rates per unit leaf area (NAR) and per unit plant mass (RGR), both measured on the basis of above-ground mass, increased with the level of defoliation at high light but not at low light. We estimated that the increases in f(lam) and NAR enabled C. elegans to compensate for part of the potential loss in performance caused by defoliation, even in cases where their RGR values were lower than those of control plants. Sensitivity analysis indicated that changes in NAR contributed more to this compensation than f(lam), but the importance of f(lam) increased with defoliation level and with decreasing light availability. The degree of compensation was higher in the high- than in the low-light treatment, suggesting that the possession of traits associated with compensatory growth. may be more important in sunny than in shaded environments. The degree of compensation differed depending on the measure of performance. Defoliated plants fully compensated for the potential reduction in lamina growth but compensated for <20% of estimated loss in reproductive output. Since survival of C. elegans plants appears to be. strongly associated with their total leaf area, the greater compensation for lamina growth is important in relation to population dynamics.
Abstract: Evolutionary variation for accumulation of small heat shock protein (sHsp) may contribute to thermal niche differentiation between species. Here we examine temperature and time-course-dependent variation for sHsp accumulation in a recently diverged pair of Encelia raised in a common environment: Encelia farinosa, common in the Mojave desert, and Encelia californica, which is found along the cool coastal bluffs of southern North America. Both species exhibit peak sHsp accumulation at 42degreesC. Encelia californica accumulated greater levels of sHsp at temperatures below 42degreesC, while E. farinosa had greater levels above 42degreesC. Encelia farinosa accumulates sHsp at temperatures up to 45degreesC, while E. californica does not synthesize sHsp above 44degreesC. Both species accumulated significant levels of sHsp while maintaining photosynthetic electron transport (F-v/F-m), but above the temperatures that elicited peak sHsp expression, levels of sHsp and F-v/F-m declined in parallel to zero. Encelia californica accumulated greater levels of sHsp more rapidly than E. farinosa following a 15 min, 42degreesC heat treatment; however, E. farinosa maintained greater F-v/F-m at all time points. Our results indicate that there are significant differences between Encelia species for sHsp accumulation but that these results depend on the duration, magnitude, and recovery time following temperature stress.
Abstract: The nuclear DNA content of angiosperms varies by several orders of magnitude. previous studies suggest that variation in 2C DNA content (i.e. the amount of DNA in G1 phase nuclei, also referred to as the 2C-value) is Correlated with environmental factors, but there are conflicting reports in the literature concerning the nature of these relationship,. We examined variation in 2C DNA content for 401 species in the ecologically diverse California flora in relation to the mean July maximum temperature, January minimum temperature, and annual precipitation within the geographical ranges of these species. Species with small 2C-values predominate in all environments. Species with large 2C-values occur at intermediate July maximum temperatures, and decline in frequency at both extremes of the July temperature gradient, and with decreasing annual precipitation. Our analysis demonstrates the utility of quantile regression for statistical inference of complex distributions such as these. The method supports our observation that relationships between nuclear DNA content and environmental factors are stronger for species with large 2C-values.
Abstract: As better phylogenetic hypotheses become available for many groups of organisms, studies in community ecology can be informed by knowledge of the evolutionary relationships among coexisting species. We note three primary approaches to integrating phylogenetic information into studies of community organization: 1. examining the phylogenetic structure of community assemblages, 2. exploring the phylogenetic basis of community niche structure, and 3. adding a community context to studies of trait evolution and biogeography. We recognize a common pattern of phylogenetic conservatism in ecological character and highlight the challenges of using phylogenies of partial lineages. We also review phylogenetic approaches to three emergent properties of communities: species diversity, relative abundance distributions, and range sizes. Methodological advances in phylogenetic supertree construction, character reconstruction, null models for community assembly and character evolution, and metrics of community phylogenetic structure underlie the recent progress in these areas. We highlight the potential for community ecologists to benefit from phylogenetic knowledge and suggest several avenues for future research.
Abstract: We examined variation in leaf size and specific leaf area (SLA) in relation to the distribution of 22 chaparral shrub species on small-scale gradients of aspect and elevation. Potential incident solar radiation (insolation) was estimated from a geographic information system to quantify microclimate affinities of these species across north- and south-facing slopes. At the community level, leaf size and SLA both declined with increasing insolation, based on average trait values for the species found in plots along the gradient. However, leaf size and SLA were not significantly correlated across species, suggesting that these two traits are decoupled and associated with different aspects of performance along this environmental gradient. For individual species, SLA was negatively correlated with species distributions along the insolation gradient, and was significantly lower in evergreen versus deciduous species. Leaf size exhibited a negative but non-significant trend in relation to insolation distribution of individual species. At the community level, variance in leaf size increased with increasing insolation. For individual species, there was a greater range of leaf size on south-facing slopes, while there was an absence of small-leaved species on north-facing slopes. These results demonstrate that analyses of plant functional traits along environmental gradients based on community level averages may obscure important aspects of trait variation and distribution among the constituent species.
Abstract: The hypothesis that species inhabiting warmer regions have greater photosynthetic tolerance of high temperatures was tested using the temperature-dependent increase in fluorescence (T-F-o). Congeneric species pairs of Atriplex, Salvia, Encelia, and Eriogonum with desert versus coastal distributions were studied in a common environment and in the field. In addition, 21 species with contrasting microclimate distributions were studied at a field site in a northern California chaparral community. The average July maximum temperature within the current distributions of species was quantified using a geographic information system. Four parameters (T-crit, T-S20, T-50, and T-max) of the T-F-o response were used to quantify photosynthetic thermotolerance. In the common environment, only the desert Atriplex species was significantly greater for all T-F,, parameters when compared to its coastal congener. In the field, desert species had significantly greater T-crit, T-S20, T-50, and T-max when compared to coastal species. The magnitude of variation between species and between genera was similar in the common environment and the field. However, T-crit, T-S20, T-50 and T-max were all significantly greater when measured in the field. There was no relationship between T-F-o parameters and the microcliniate distribution of the 21 species at the chaparral field site. In addition, T-F-o parameters for all 35 species were not correlated with the average July maximum temperature within the species ranges. However, there was a significant negative correlation between the average annual amount of precipitation inside species’ ranges and T-S20. Our results show that photosynthetic thermotolerance is (1) significantly different between genera and species, (2) highly plastic, (3) not necessarily greater for species with warm climate distributions when measured in a common environment, but (4) significantly greater overall for desert species compared to coastal species when measured in the field.
Abstract: Fire may act as a selective force on plants both through its direct effects by killing or wounding susceptible individuals and through its effect on the environment: the post-fire environment may select specific physiological traits or life historiesWe used phylogenetic independent contrasts to test the hypothesis that fire has selected for correlated evolution among alternative suites of traits in pines: a survival/avoidance suite characterized by thick bark, height, and self-pruning of dead branches; and a fire-embracing strategy in which plants invest little into survival, exhibit traits which enhance flammability, and use fire as a means to cue seedling establishment to the post-fire environment through serotinous conesWe created a set of alternative ’supertree’ phylogenies for the genus Pinus from published sourcesUsing these alternative phylogenies, published ecological data for 38 pine species, and newly collected morphological data, we demonstrate that much variation in trait evolution occurs along a fire-surviving/fire-embracing axisPines vary in their susceptibility to ignition since a tree that retains dead branches is more likely to carry a fire into the canopy than a tree that self-prunesThe evolution of increased flammability may have altered evolutionary trajectories prompting an evolutionary switch from a fire-surviving to a fire-embracing life history. Alternatively, the fire-embracing strategy may in fact select for increased flammability to ensure canopy ignition and the realization of serotinous seed-release.
Abstract: Interspecific variation in chloroplast low molecular weight (cLMW) HSP (heat shock protein) expression was examined with respect to phylogeny, species specific leaf area, chlorophyll fluorescence, and mean environmental conditions within species ranges. Eight species of Ceanothus (Rhamnaceae) were heat shocked for 4 h at several different temperatures. Leaf samples were collected immediately after the heat shock, and cLMW HSP expression was quantified using Western blots. At 45°C species from the subgenus Cerastes had significantly greater cLMW HSP expression than species from the subgenus Ceanothus. Specific leaf area was negatively correlated with cLMW HSP expression after the 45°C heat treatment. In addition, chlorophyll fluorescence (F(v)/F(m)) 1 h after the heat shocks was positively correlated with cLMW HSP expression. Contrary to our prediction, there was no correlation between July maximum temperature within species ranges and cLMW HSP expression. These results suggest that evolutionary differentiation in cLMW HSP expression is associated with leaf physiological parameters and related aspects of life history, yet associations between climatic conditions within species ranges and cLMW HSP expression require further study.
Abstract: 1. A new method (the iterative approach) is presented by which growth analyses can be conducted on plants that have been subjected to significant losses in biomass and leaf area between harvests. The method is particularly useful to analyse the effects of defoliation on growth and biomass allocation.2. Values for the following parameters can be estimated: absolute growth rate (g day(-1)), relative growth rate (RGR, g g(-1) day(-1)), net assimilation rate (NAR, g m(-2) day(-1)), leaf area ratio (LAR. m(2) g(-1)). fraction of newly assimilated mass that is allocated to leaf lamina production (f(lam), g g(-1)), and daily fractional change in the average specific leaf area of plants (rho, day(-1)). These parameters are determined by means of iterations. We defined a number of growth functions. and the values of NAR,f(lam) and the SLA of newly produced leaves were changed until these functions correctly predicted the measured total plant mass. leaf lamina mass and leaf area at the end of the growth period. This avoids having to assume a constant relationship between leaf area and biomass (as in the ’classical’ approach), and it avoids the use of polynomial functions to fit growth data (as in the ’functional’ approach) that are unsuitable for fitting data sets exhibiting discontinuities such as abrupt changes in biomass.3. The method was applied to a greenhouse experiment in which we analysed the effects of sustained defoliation on growth and biomass allocation in a tropical understorey palm, Chamaedorea elegans Mart.4. We showed that C elegans plants respond to defoliation with a considerable increase in the allocation of new assimilates to lamina growth (f(lam)) and that, despite the repeated loss of leaf area and associated reductions in LAR, they had RGR values that were similar to those of undamaged plants.
Abstract: Independent contrasts are widely used to incorporate phylogenetic information into studies of continuous traits, particularly analyses of evolutionary trait correlations, but the effects of taxon sampling on these analyses have received little attention. In this paper, simulations were used to investigate the effects of taxon sampling patterns and alternative branch length assignments on the statistical performance of correlation coefficients and sign tests; "full-tree" analyses based on contrasts at all nodes and "paired-comparisons" based only on contrasts of terminal taxon pairs were also compared. The simulations showed that random samples, with respect to the traits under consideration, provide statistically robust estimates of trait correlations. However, exact significance tests are highly dependent on appropriate branch length information; equal branch lengths maintain lower Type I error than alternative topological approaches, and adjusted critical values of the independent contrast correlation coefficient are provided for use with equal branch lengths. Nonrandom samples, with respect to univariate or bivariate trait distributions, introduce discrepancies between interspecific and phylogenetically structured analyses and bias estimates of underlying evolutionary correlations. Examples of nonrandom sampling processes may include community assembly processes, convergent evolution under local adaptive pressures, selection of a nonrandom sample of species from a habitat or life-history group, or investigator bias. Correlation analyses based on species pairs comparisons, while ignoring deeper relationships, entail significant loss of statistical power and as a result provide a conservative test of trait associations. Paired comparisons in which species differ by a large amount in one trait, a method introduced in comparative plant ecology, have appropriate Type I error rates and high statistical power, but do not correctly estimate the magnitude of trait correlations. Sign tests, based on full-tree or paired-comparison approaches, are highly reliable across a wide range of sampling scenarios, in terms of Type I error rates, but have very low power. These results provide guidance for selecting species and applying comparative methods to optimize the performance of statistical tests of trait associations.
Abstract: We developed a description of a central New England deciduous hardwood forest based on the distribution within the community of morphological and life history traits (N = 34) and environmental factors. Classification by TWINSPAN of 186 species based on morphological and life history traits identified six major functional groups of species largely corresponding to growth form. A data matrix of plots x traits was ordinated using PCA. Each of the resulting four PCA axes was associated with a major environmental gradient: drainage, site exposure, disturbance due to past land use and degree of disturbance in the 1938 New England Hurricane (24.9, 19.8, 11.7 and 8.4% of the variation respectively). Two patterns suggested that a suite of potentially functional traits, rather than a few key characters (e.g. vital attributes), govern the distribution of species in this community: (1) each of the four axes was largely associated with a different group of traits and (2) each axis was associated with several traits that appeared to sort independently (i.e. not to co-occur within species). Evaluating one often-examined trait, there was no evidence that dispersal ability limited the colonization of species into secondary woodlands. We also found that landscape-scale abundance was associated with a small number of traits. Production of fleshy fruits and few diaspores per plant were positively associated with landscape-level abundance. Our results suggest that attempts to understand the overall structure and function of this plant community based on a few key characters, such as dispersal ability, will meet with limited success. However, when focusing on one aspect of the community, such as frequency across the landscape, relatively few characters may be important.
Abstract: Prior studies of a broad array of seed plants have reported strong correlations among leaf life span, specific leaf area, nitrogen concentration, and carbon assimilation rates, which have been interpreted as evidence of coordinated leaf physiological strategies. However, it is not known whether these relationships reflect patterns of evolutionary convergence, or whether they are due to contrasting characteristics of major seed plant lineages. We reevaluated a published data set for these seven traits measured in over 100 species, using phylogenetic independent contrasts calculated over a range of alternative seed plant phylogenies derived from recent molecular systematic analyses. In general, pairwise correlations among these seven traits were similar with and without consideration of phylogeny, and results were robust over a range of alternative phylogenies. We also evaluated relationships between these seven traits and lamina area, another important aspect of leaf function, and found moderate correlations with specific leaf area (0.64), mass-based photosynthesis (0.54), area-based nitrogen (-0.56), and leaf life span (-0.42). However, several of these correlations were markedly reduced using independent contrasts; for example, the correlation between leaf life span and lamina area was reduced to close to zero. This change reflects the large differences in both these traits between conifers and angiosperms and the absence of a relationship between the traits within these groups. This analysis illustrates that most interspecific relationships among leaf functional traits, considered across a broad range of seed plant taxa, reflect significant patterns of correlated evolutionary change, lending further support to the adaptive interpretation of these relationships.
Abstract: A simple model of shoot-level carbon gain is presented addressing the optimal number and life span of leaves in relation to alternative optimality criteria: (1) maximizing carbon export from the shoot, or (?) maximizing the rate of leaf production at the shoot tip. Additionally, the processes that cause declining assimilation with leaf age are considered in relation to (1) leaf position on the shoot (e.g., self-shading) versus (2) leaf age per se. Using these alternative scenarios, only a model based on position-dependent assimilation and maximization of leaf production rates resulted in quantitative predictions for all aspects of leaf dynamics on the shoot (i.e., leaf number, life span, and birth rate), while other approaches predicted that one or more parameters would be infinite. This formulation of the model also predicted that leaves should be maintained on the shoot until the diurnal carbon balance declines to zero, in contrast with other scenarios which predict that leaves should be shed while maintaining a positive carbon balance. Predictions of the model were supported by the results of a field study of carbon gain and leaf dynamics in saplings of three species of tropical pioneer trees (Carica papaya, Cecropia obtusifolia, and Hampea nutricia) which differ in the number of leaves per shoot. The results illustrate that in these fast-growing plants, leaf production and height growth may be more appropriate measures of performance than net carbon export from the shoot, and suggest that leaf senescence is primarily a function of the position of a leaf within the canopy, rather than its chronological age.
Abstract: We studied the evolution of leaf size, sapling canopy allometry, and related traits in 17 Acer species growing in the understory of temperate deciduous forests, using parsimony methods, randomization tests, and independent contrasts calculated on a phylogeny inferred from nuclear ribosomal internal transcribed spacer (ITS) sequences. Bivariate correlations and multivariate analyses indicated two independent suites of coevolving traits, and the results were robust over a range of alternative phylogenies. The first suite consisted of strong positive correlations among twig thickness, leaf size, inflorescence length, and branch spacing (Corner's rules). Seed size and mature height were also weakly correlated with these traits. The second suite reflected aspects of sapling crown allometry, including crown size, stem diameter, and total leaf area, which appear to be related to shade tolerance. There was a weak negative correlation between sapling crown size and mature height, but no correlation with leaf or seed size. Most correlations were similar in magnitude for ahistorical and independent contrasts analyses, and discrepancies between these two measures were greater in traits with lower levels of convergent evolution. The evolutionary correlations among twig, leaf, seed, inflorescence, and canopy dimensions emphasize the need for integrated theories of evolution and function of these disparate traits.
Abstract: We studied the effects of CO2 elevation on leaf and canopy photosynthesis and optimal leaf area index (LAI) for stands of the annual species Abutilon theophrasti and Ambrosia artemisiifolia. Leaf photosynthesis was modeled as a function of photosynthetic photon flux density (PPFD) and nitrogen content per unit leaf area (N-L). There was a curvilinear relationship between the light-saturated rates of leaf photosynthesis (P-max) and N-L. CO2 elevation significantly increased P-max as a function of N, in both species. Dark respiration (R-d) was linearly correlated with N-L. CO2 elevation slightly but significantly increased R-d in Abutilon, while it had no significant effect on R-d in Ambrosia. The initial slope of a light-response curve was determined from quantum yield (phi(abs)) multiplied by leaf absorptance and then calibrated against N-L. Daily canopy photosynthesis, calculated by integration of leaf photosynthesis with the actual distribution of leaf area, leaf N, and PPFD within a canopy, showed fairly good agreement with the canopy photosynthesis estimated from growth analysis. CO2 elevation increased canopy photosynthesis by 30-50%. Based on the leaf photosynthesis model for Abutilon, we calculated daily canopy photosynthesis for a given LAI and N availability, in which N was assumed to be distributed optimally within a leaf canopy to maximize daily canopy photosynthesis. An optimal LAI to maximize daily canopy photosynthesis was obtained for each level of N availability and this optimum increased with increasing N availability. Contrary to the often predicted increase in LAI with CO2 elevation, the optimum LAI did not increase at high CO2 when N availability was limited. Two factors were suggested to be involved in counteracting the increase in LAI in a high-CO2 world. One is the higher phi(abs) of plants grown in elevated CO2, which makes leaves in the canopy more N limited, favors higher N-L and thus lowers optimal LAI. The other is the higher R-d in elevated CO2, which leads to higher light compensation points, and lowers optimal LAI.
Abstract: Phylogenetic comparative analyses combine information on character states and phylogenetic relationships of taxa to test hypotheses regarding character evolution. These studies encounter uncertainties at various steps, including uncertainty in the topology of phylogenetic trees, the scoring of characters, and the addition of taxa that have not explicitly been included in phylogenetic analyses. Here we highlight a variety of sensitivity tests designed to explore the robustness of comparative conclusions to changes in underlying assumptions. These include the examination of character correlations on a set of plausible phylogenetic hypotheses (including alternative rootings and ‘neighbouring’ trees), as well as under alternative character codings. TreeBASE - a prototype relational database of phylogenetic data - should prove useful in accessing alternative hypotheses.
Abstract: In forest gaps, the geometry of canopy openings above a plant determines the spatial distribution of diffuse and direct radiation, which may be received from different sectors of the sky. We examined crown orientation and light interception by seedlings of four species of pioneer trees, Cecropia obtusifolia, Heliocarpus appendiculatus, Piper auritum, and Trema micrantha, transplanted into natural forest gaps at the Los Tuxtlas Tropical Biology Station, Veracruz, Mexico. The four species differed in the number and mean size of leaves, but total leaf area was not significantly different among equal age plants. The mean azimuth and zenith angle of diffuse and direct radiation were determined from hemispherical photographs taken above each plant. Mean diffuse radiation vectors were distributed around the zenith, with a significant clustering in an easterly direction, while mean direct radiation vectors were distributed along the solar track in the southern portion of the sky. The mean orientation of the crown of each seedling was determined from detailed, three-dimensional reconstruction of leaf display using a canopy architecture model. A directional correlation test indicated that seedling crowns were oriented toward diffuse radiation received from the gap, and not toward direct radiation. The mean orientation of the crown, at the whole plant level, was primarily determined by nonrandom orientation of individual leaves. The mechanisms of this differential response to diffuse vs. direct radiation are not known. For diffuse radiation, the total light capture capacity integrated over all sky directions increased with seedling leaf area, while efficiency of capture per unit area decreased. Based on simulations, the light interception efficiency of each plant in its own microsite was 5-25% greater than would be achieved in the sites of conspecifics or in symmetric canopy openings centered on the zenith. These analyses provide a quantitative estimate of the value of nonrandom leaf and crown orientation for whole-plant light interception in natural environments.
Abstract: The effects of rising atmospheric CO2 concentrations on natural plant communities will depend upon the cumulative responses of plant growth and reproduction to gradual, incremental changes in climatic conditions. We analysed published studies of plant responses to elevated CO2 to address whether reproductive and total biomass exhibit similar enhancement to elevated vs. ambient CO2 concentrations, and to assess the patterns of plant response along gradients of CO2 concentrations. In six annual plant species, mean enhancement at double ambient vs. ambient CO2 was 1.13 for total biomass and 1.30 for reproductive biomass. The two measures were significantly correlated, but there was considerable scatter in the relationship, indicating that reproductive responses cannot be consistently predicted from enhancement of total biomass. Along experimental CO2 gradients utilizing three concentrations, there was a great diversity of response patterns, including positive, negative, non-monotonic and non-significant (nat) responses. The distribution of response patterns differed for plants grown in stands compared to those grown individually. Positive responses were less frequent in competitive environments, and non-monotonic responses were more frequent. These results emphasize that interpolation of plant response based on enhancement ratios measured at elevated vs. ambient CO2 concentrations is not sufficient to predict community responses to incremental changes in atmospheric conditions. The consequences of differential response patterns were assessed in a simulation of community dynamics for four species of annual plants. The model illustrates that the final community composition at a future point in time depends critically on both the magnitude and the rate of increase of atmospheric CO2.
Abstract: We used path analysis to ask whether leaf position or leaf light level was a better predictor of within-plant variation in leaf nitrogen concentration in five species of rain forest pioneer trees (Cecropia obtusifolia, Ficus insipida, Heliocarpus appendiculatus, Piper auritum, and Urera caracasana) from the Los Tuxtlas Biological Station, Veracruz, Mexico. Three hundred seventy-five leaves on 28 plants of the five species were analyzed for leaf nitrogen concentration, leaf mass per area, and leaf light interception at different positions (= nodes) along a shoot. Mean values of leaf nitrogen concentration ranged from 0.697 to 0.993 g/m(2) in the five species, and varied by as much as 2.24 g/m(2) among leaves on individual plants. Leaf position on the shoot explained significantly more of the within-plant variation in leaf nitrogen concentration than did leaf light level in four of the five species: Cecropia obtusifolia, Heliocarpus appendiculatus, Piper auritum (branch leaves only), and Urera caracasana. However, individual species differed considerably in the patterns of nitrogen allocation and leaf mass per area among leaves on a shoot. These results suggest that leaf nitrogen deployment in these plants is, in part, developmentally constrained and related to the predictability of canopy light distribution associated with plant growth form.
Abstract: Plant ecologists and evolutionary biologists frequently examine patterns of phenotypic variation across variable environments or genetic identities. Too often, we ignore the fact that most phenotypic traits change throughout growth and development of individual plants, and that rates of growth and development are highly variable. Plants growing in different environments are likely to grow at different rates, and will be of different sizes and stages of development at a particular age. When we compare plants as a function of plant size or developmental stage, as well as a function of age, we broaden our understanding of phenotypic variation between plants.
Abstract: Tropical vines in the Araceae family commonly exhibit alternating periods of upward and downward growth, decoupling the usual relationship between decreasing light environment with increasing age among the leaves on a shoot. In this study I examined patterns of light, leaf specific mass, and leaf nitrogen concentration in relation to leaf position, a measure of developmental age, in field collected shoots of Syngonium podophyllum. These data were analyzed to test the hypothesis that nitrogen allocation parallels within-shoot gradients of light availability, regardless of the relationship between light and leaf age. I found that leaf nitrogen concentration, on a mass basis, was weakly correlated with leaf level light environment. However, leaf specific mass, and consequently nitrogen per unit leaf area, were positively correlated with gradients of light within the shoot, and either increased or decreased with leaf age, providing support for the hypothesis that nitrogen allocation parallels gradients of light availability.
Abstract: 1. Populations of an annual plant, Abutilon theophrasti, were grown at four densities (100, 500, 1500 and 4000 m-2) and two CO2, concentrations (350 and 700 mul l-1) to examine the influence of CO2 environment on density-dependent patterns of demography and reproduction. Variables measured included survivorship, proportion of plants flowering and fruiting, number of fruiting individuals, number of seeds per individual, total seed production per population, mean seed mass, and germination of seeds produced in each environment.2. All variables, except the number of fruiting individuals, declined with increasing density, and at the highest density no individuals set seed. The number of fruiting individuals was highest at a density of 500m-2. In the elevated CO2 environment, survivorship was significantly reduced but the proportion of plants flowering and fruiting and the number of fruiting individuals in each population all increased. Total population seed production was higher in the elevated CO2 environment at all densities, although the differences were not significant. Significant effects of CO2, concentration were observed only for population-level variables, but not for mean individual fecundity or seed size. Seed germination declined with increasing maternal density, and no germination was recorded for seeds produced at 1500 m-2 3. Simple models of population dynamics, utilizing difference equations, were constructed to examine potential population-level consequences of these density and CO2 effects. In the absence of a persistent seed pool, the simulated populations exhibited damped or stable oscillations under low germination values, but displayed non-cyclic (’chaotic’) oscillations or went extinct for higher germination due to the complete failure of seed-set at high density. Because of its higher fecundity, the elevated-CO2 population generally exhibited greater oscillations, and the critical germination value at which the simulated populations went extinct was much lower for the elevated-CO2 than for the ambient-CO2 population.
