Phylogenetic structure of Brazilian savannas under different fire regimes

Questions: Fire is a strong filter in fire‐prone communities and is expected to assemble closely related species when functional traits are conserved in plant lineages. Do frequent fires assemble savannas with closely related species (phylogenetic clustering)? If so, what are the clades pruned by fire in the phylogenetic trees? Are species of semi‐deciduous seasonal forests, where fires are not frequent, less related than expected by chance (phylogenetic over‐dispersion)? Are life forms conserved in the phylogeny of the species? Location: Central and SE Brazilian savannas (Emas National Park, 18°18′S, 52°54′W; Brasília, 15°56′–15°57′S, 47°53′–47°56′W and Corumbataí‐Itirapina, 22°13′–22°15′S, 47°37′–47°39′W); and close semi‐deciduous seasonal forests (in Pirenópolis, 15°45′S, 49°04′W; Brasília, 15°33′S, 47°51′W; and São Carlos, 21°55′S, 47°48′W). Methods: We recorded woody species in savannas under different fire regimes and in semi‐deciduous seasonal forests. We obtained data from the literature and from field sampling. We compared mean phylogenetic distance of species of savanna and of nearby semi‐deciduous seasonal forest sites. We obtained significance by randomizing the species among the tips of phylogenetic trees. We also assessed whether life forms were evolutionary conserved across phylogeny of the studied plants (phylogenetic signal) with tests based on the variance of phylogenetic independent contrasts. Results: Some sites of savanna under high fire frequency were characterized by phylogenetic over‐dispersion of woody species whereas, in contrast, some sites of semi‐deciduous seasonal forest were characterized by phylogenetic clustering. We found phylogenetic signals in the traits across the phylogeny of the 801 species investigated. Conclusion: Fire may have different roles in assembling plant species in Brazilian savannas than in other fire‐prone communities. We postulate that the absence of phylogenetic clustering in the cerrado is mainly due to the persistence of long‐lived resprouting species from different plant lineages.     

Latitudinal clines in body size,but not in thermal tolerance or heat‐shock cognate 70 (HSC70), in the highly‐dispersing intertidal gastropod Littorina keenae (Gastropoda: Littorinidae)

Natural populations of widely‐distributed animals often exhibit clinal variation in phenotypic traits or in allele frequencies of a particular gene over their geographical range. A planktotrophic intertidal snail, Littorina keenae is broadly distributed along the north‐eastern Pacific coast through a large latitudinal range (24°50′N–43°18′N). We tested for latitudinal clines in two complex phenotypic traits – thermal tolerance and body size – and one single locus trait – heat shock cognate 70 (HSC70) – in L. keenae along almost its entire geographical range. We found only weak evidence for a latitudinal cline in the thermal tolerance and no evidence for a cline in allele frequencies at HSC70. However, as predicted by Bergmann's rule, we detected a strong latitudinal cline that accounted for 60% of the variance in body size (R² = 0.598; P < 0.001). In contrast, body size did not significantly affect thermal tolerance. HSC70 showed no genetic differentiation among the populations, supporting our previous mitochondrial gene‐based estimate of high gene flow during this snail's free‐swimming larval stage. Given that L. keenae experiences panmixia along its species range, the observed size cline may be partially or entirely caused by a phenotypically plastic response to local thermal environments rather than by genetic divergence in body size among populations in response to locally optimizing natural selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 494–505.     

The importance of spatial scale for trait–abundance relations

The concept of community assembly through trait‐based environmental filtering has played a key role in our understanding of how communities change over space and time, however, the importance of spatial scale in the filtering process remains unclear. We propose that different environmental filters may operate at different spatial scales, and that filters at finer scales would be nested within those acting at coarser scales. We tested for the existence of spatially nested sets of trait‐based filters in a temperate native grassland by applying the recently proposed maximum entropy (MaxEnt) approach to trait‐based community assembly, which we extend through a trait selection procedure. We found that different traits were important in influencing the abundances of species at the three different spatial scales examined (micro‐habitat, habitat, landscape), supporting the idea that trait based filtering processes operating at coarse spatial scales can be quite distinct from those operating at fine scales. Despite this result, we identified several traits which were frequently related to abundance at all spatial scales. Taken together, our results support the proposition that trait‐based environmental filters at finer spatial scales are nested within those operating at coarser scales. We compared our results to those obtained using a simpler trait‐by‐trait analytical approach (correlation analysis and MaxEnt on individual traits). The capacity for MaxEnt to incorporate multiple traits simultaneously provided unique insights into the important traits at each spatial scale and presents significant advantages over existing univariate and multivariate approaches.     

Traits of benthic macroinvertebrates in semi-natural French streams: an initial application to biomonitoring in Europe

• 1 The methods used to indicate the biological state of streams are often based on taxonomic composition, and the abundance of species or other taxa. This ‘taxonomic structure’ varies among ecoregions and cannot be applied to wider geographical areas. Therefore, we assessed the species traits of benthic macroinvertebrates from semi‐natural reference sites as a potential benchmark for large‐scale biomonitoring. Our purpose was to assess the stability of community structure, based on the representation of taxa and of traits, across large gradients of geology (sedimentary to granitic), altitude (65–1982 m), geographical coordinates (0° 48′ W to 7° 20′ E and 42° 52′ to 48° 44′ N), stream order (1–5) and slope (0.5–60‰).
• 2 We used invertebrate abundance data from the 62 most natural French stream sites available. These abundance data served to weight the occurrence of ‘biological’ traits, such as reproductive characteristics, mobility, resistance forms, food, feeding habits, respiration, and ‘ecological’ traits, such as preferences for temperature, trophic level, saprobity, biogeographic distribution, longitudinal zonation, substratum and current velocity.
• 3 Multivariate analyses of taxonomic composition demonstrated a clear site gradient from lowlands to uplands and from calcareous to granitic geology. In contrast, community structure based on both biological and ecological traits was stable across environmental gradients.
• 4 The frequency distribution of biological traits indicated that the stream benthos of the ‘reference sites’ had a mixture of categories which confirmed theoretical predictions for temporally stable and spatially variable habitats. A mixture of ecological trait categories also occurred at our reference sites. Thus, semi‐natural benthic macroinvertebrate communities are functionally diverse. Moreover, we included an initial application of these traits to a case of slightly to moderately polluted sites to show that the impact of humans significantly changes this natural functional diversity.
• 5 Future studies should focus on the potential for various biological and ecological traits to discriminate different human impacts on the benthic macroinvertebrates of running waters, and on the integration of this functional application into a general ‘reference‐condition’ approach.

     

Plant species traits across a riparian‐zone/forest ecotone

Abstract. We examined the changes in prevalence of nine plant traits – including the presence of woody stem tissue, leaf longevity, nitrogen fixation, seed longevity, dispersal vector, pollination vector, and clonal growth form – across a riparian/forest‐understory ecotone. This ecotone, found along headwater streams in boreal mixed‐wood forests, supports four distinct vegetation zones: streambank, riparian, transition, and upland forest understory. The objective of this study was to identify specific trait patterns that may indicate functional responses to the changes in environmental factors such as nutrient availability and wind exposure that occur across the ecotone. The suites of plant species traits found in each zone were distinct, with a strong change in the prevalence of several traits. Wind and insect pollination, wind and vertebrate diaspore dispersal, and deciduous and evergreen leaves showed the greatest change in prevalence between the vegetation types. Some traits, including insect pollination and vertebrate diaspore dispersal, were strongly correlated within species. The consistent cooccurrence of pairs of traits in the same species suggests common responses by very different traits to the same environmental factor. This study demonstrates that an ecotone can be characterized not only as a discontinuity in species distributions or environmental factors, but also as a discontinuity in the trait spectrum. Examining ecotones from a trait perspective has strong potential for identifying the environmental factors and associated species functional responses that encourage the development of distinct vegetation boundaries.     

Evaluating life‐history strategies of reef corals from species traits

Classifying the biological traits of organisms can test conceptual frameworks of life‐history strategies and allow for predictions of how different species may respond to environmental disturbances. We apply a trait‐based classification approach to a complex and threatened group of species, scleractinian corals. Using hierarchical clustering and random forests analyses, we identify up to four life‐history strategies that appear globally consistent across 143 species of reef corals: competitive, weedy, stress‐tolerant and generalist taxa, which are primarily separated by colony morphology, growth rate and reproductive mode. Documented shifts towards stress‐tolerant, generalist and weedy species in coral reef communities are consistent with the expected responses of these life‐history strategies. Our quantitative trait‐based approach to classifying life‐history strategies is objective, applicable to any taxa and a powerful tool that can be used to evaluate theories of community ecology and predict the impact of environmental and anthropogenic stressors on species assemblages.     

Thermal biology of mosquito‐borne disease

Mosquito‐borne diseases cause a major burden of disease worldwide. The vital rates of these ectothermic vectors and parasites respond strongly and nonlinearly to temperature and therefore to climate change. Here, we review how trait‐based approaches can synthesise and mechanistically predict the temperature dependence of transmission across vectors, pathogens, and environments. We present 11 pathogens transmitted by 15 different mosquito species – including globally important diseases like malaria, dengue, and Zika – synthesised from previously published studies. Transmission varied strongly and unimodally with temperature, peaking at 23–29ºC and declining to zero below 9–23ºC and above 32–38ºC. Different traits restricted transmission at low versus high temperatures, and temperature effects on transmission varied by both mosquito and parasite species. Temperate pathogens exhibit broader thermal ranges and cooler thermal minima and optima than tropical pathogens. Among tropical pathogens, malaria and Ross River virus had lower thermal optima (25–26ºC) while dengue and Zika viruses had the highest (29ºC) thermal optima. We expect warming to increase transmission below thermal optima but decrease transmission above optima. Key directions for future work include linking mechanistic models to field transmission, combining temperature effects with control measures, incorporating trait variation and temperature variation, and investigating climate adaptation and migration.     

Mycorrhizal traits and plant communities: perspectives for integration

Plant functional type‐ and trait‐based approaches to understanding vegetation dynamics are gradually gaining popularity. However, plant mycorrhizal traits are rarely considered in plant trait databases and are almost totally neglected in trait‐based plant community studies, despite more than 90% of the land flora being mycorrhizal. In this paper I describe and define the mycorrhizal traits of plant species, notably mycorrhizal type, mycorrhizal status, mycorrhizal flexibility and mycorrhizal dependency, which potentially influence plant distribution and community structure. I propose ways of using these traits for large‐scale synthetic studies for understanding the role of mycorrhizal symbiosis in vegetation dynamics. I suggest considering plant community mycorrhization – community means of mycorrhizal traits weighted by plant species abundances – and suggest an index of mycorrhization to describe the mycorrhizal trait composition of plant communities.     

Leaf palatability and decomposability increase during a 200‐year‐old post‐cultural woody succession in New Zealand

Question: How do aggregate trait values and functional diversity of leaf traits linked to palatability and decomposability change during a woody post‐cultural succession spanning 200 years? Location: Coastal Marlborough, South Island, New Zealand. Methods: The biomass of all woody species was determined in 32 20‐m × 20‐m plots ranging from 10 to 200 years in time since last disturbance. Species abundances were combined with data on leaf nutrient, secondary metabolite and structural carbohydrate content to calculate biomass‐weighted trait means (i.e. aggregate trait values) and functional diversity index values for each plot. Aggregate trait values and functional diversity were regressed on successional age and total live above‐ground carbon content to examine functional shifts with succession and one consequence of succession – increasing above‐ground carbon. Results: Almost all significant regressions between aggregate trait values and both successional age and above‐ground carbon indicated a shift toward increased leaf palatability and decomposability during succession. The relationships were all non‐linear, with aggregate trait value shifts occurring relatively early in the successional sequence. There was weak evidence for an increase in functional richness with succession, but this was a secondary effect relative to the shifts in aggregate trait values. Conclusions: These results are in direct contrast with studies of the early stages of herbaceous post‐cultural successions from grasslands to shrublands, which have found a shift towards communities of decreasing palatability and decomposability, suggesting that functional shifts in woody succession may be fundamentally different.     

Testing for the influence of niche and neutral factors on sapling community assembly beneath isolated woody plants in grasslands

Questions: Perching and nursing effects drive initial steps of forest expansion over grasslands. Nursing effect is obviously related to niche mechanisms, while perching effect is likely to result both from neutral and niche factors. This study assessed the effect of neutral and niche factors on species composition in sapling communities developing beneath isolated trees/shrubs (ITS) in grassland. Location: A mosaic of Campos grassland and Araucaria forest in São Francisco de Paula, southern Brazil (29°28′S, 50°13′W). Methods: We described sapling communities beneath 32 ITS using mean number of forest woody saplings of different species. We performed a stepwise canonical correspondence analysis (CCA) to select ITS traits that maximized the association with species composition. Then we evaluated the contribution of distance from seed source, ITS traits and distance‐structured ITS traits on sapling community assembly using a variation partitioning method based on CCA and partial CCA. Results: Sapling species composition was significantly explained by ITS traits (ITS dispersal mode, ITS growth form, crown area:ITS height ratio, crown area, ITS height and crown area:volume ratio). Distance from seed sources explained only a minor, non‐significant fraction of sapling species composition. Distance‐structured trait variation was negligible. Conclusions: Sapling community assembly beneath ITS was mostly explained by niche factors related to both nursing and perching effects. Dispersal limitation explained only a small fraction of variation in species composition beneath ITS, suggesting that neutral‐based perching effect had a minor role in community assembly.     

Contrasting patterns of trait‐based community assembly in lianas and trees from temperate Australia

Trait variation in plant communities is thought to be constrained by two opposing community assembly processes operating at discrete spatial scales: habitat filtering and limiting similarity between coexisting species. Filtering processes cause convergence in ecological strategy as species are excluded from unsuitable sites, whilst limiting similarity leads to the divergence of trait values between co‐occurring species in order to alleviate competition for finite resources. Levels of alpha (within‐site) and beta (among‐site) trait variation can be indicative of the strength of these community assembly processes. We used trait‐gradient analysis to explicitly compare evidence of community assembly patterns in lianas (woody vines) and trees. These two growth forms exhibit striking differences in carbon capture and regeneration strategies, yet trait‐based mechanisms that maintain their coexistence remain understudied. Using data for four functional traits – leaf mass per area, leaf nitrogen content (N_mass), leaf area and seed mass – we partitioned interspecific trait variation in lianas and trees into alpha and beta components. Our three key findings were: 1) lianas and trees exhibit divergent patterns of trait‐based habitat filtering, due to differences in the relationship between leaf size and the other three traits examined (LMA, N_mass and seed mass), 2) on average, liana species possess smaller seeds, lower LMA and higher N_mass than do trees, but there was no clear difference in leaf area between the two growth forms, and 3) soil fertility was correlated with trait variation (leaf area, seed mass) at the site‐level in trees, but not in lianas. These results provide evidence that dominant growth forms can be filtered into the same habitat on the basis of different combinations of traits. Our findings have important implications for community assembly and co‐existence theory and for more pragmatic matters such as using trait‐based principles to inform community restoration.     

A test of ecological succession hypotheses using 55‐year time‐series data for 361 boreal forest stands

Aim There has been much work on succession over many decades, but succession fundamentals are still debated because of the reliance on chronosequences and dendrochronological reconstruction, both of which are problematic approaches. Here we use time‐series data to test four hypotheses that lie at the heart of successional theory: (1) the neighbourhood effect hypothesis – tree species abundance is time dependent; (2) the density‐dependence hypothesis – a rare species is more favoured over time; (3) the resource ratio hypothesis – species that can grow at the lowest resource level tend to dominate resource limited sites through succession; and (4) the intermediate disturbance hypothesis – intermediate disturbances increase the abundance of rare species. Location Central boreal forest in Canada (47°50′–50°10′ N; 80°10′–85°50′ W). Methods We used repeated measurements from sequential aerial photography and ground surveys for 361 fire‐origin stands that were measured over a c. 55‐year period. Results Shade‐intolerant Pinus banksiana decreased, tolerant Thuja occidentalis increased, intolerant Populus spp. and Betula papyrifera displayed a U‐shaped trend, and intermediate‐tolerant Picea spp. and tolerant Abies balsamea did not change with time since fire, showing evidence of negative, positive, or neutral neighbourhood effects. Species either persisted for longer or increased more in non‐conspecific stands, and had higher increases in abundance when associated with species of contrasting shade tolerance, supporting the density‐dependence hypothesis and indicating shade‐tolerance complementarity as a mechanism for coexistence. Resource‐poor soils favoured those species capable of tolerating limited resources, whereas rich soils permitted invasion and promoted mixtures, supporting the resource ratio hypothesis. Intermediate disturbances increased the invasibility of rare species. Main conclusions Contrary to previous studies where time since a stand‐replacing disturbance is the sole predictor for succession, our study shows that time alone is either an insufficient predictor or is irrelevant to species dynamics in the boreal forest. Rather, density dependence, site resource and intermediate disturbances are key mechanisms in species dynamics and coexistence over time.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Fifty‐seven years of composition change in the eastern boreal forest of Canada

Question: In the boreal forest of eastern Canada, how does forest vegetation change in the sustained absence of fire? Location: Eastern boreal forest in Quebec's North Shore region, Canada (49°30′–50°00′N; 67°30′–68°35′W). Methods: Aerial photos from three different periods (1930, 1965 and 1987) were used to characterize changes in vegetation composition in 23 scenes of 200 ha. Time since fire, presence of secondary disturbances and data on soil and topographic variables were obtained. Ordination and clustering techniques were used to define compositional trajectories of change over the 57‐yr period. These trajectories were further grouped into pathways based on compositional changes, time since fire and preferential deposit‐drainage types. Results: Among the 26 compositional trajectories, three successional pathways were distinguished. Two start post‐fire succession with a dominance of intolerant hardwood. In one of these, this is followed by an increase in Abies balsamea, while in the second the importance of Picea mariana increases with time. In the third pathway P. mariana is an important component from the outset. In this pathway, we observed modest fluctuation in the relative dominance of P. mariana and A. balsamea and variation in stand structure. Conclusion: The boreal forest vegetation of Eastern Canada is diverse and dynamic even in the absence of fire, notably under the influence of partial disturbances. Such disturbances can be associated with changes in composition or stand structure. The development of management strategies aimed at maintaining stand diversity by emulating a broader variety of partial and secondary disturbances should be encouraged.     

生长型分类方案不同导致森林生态系统植物功能性状的统计偏差

将植物划分为不同的生长型来统计植物功能性状特征,是当前植物性状研究中常用的方法;但生长型分类方案的不同很可能造成植物功能性状统计分析的偏差,对此偏差的评估却尚未见报道。根据植物志描述及野外调查实际情况,将生长型划分为3种不同的分类方案：分类1：根据植物志信息划分为传统意义的乔木和灌木;分类2：根据树高和胸径划分乔木、小乔木和灌木;分类3：仅根据树高划分乔木层与灌木层的乔木和灌木。以东部亚热带常绿阔叶林区域的浙江金华北山35种优势阔叶木本植物的枝叶性状为研究对象,比较不同生长型分类对植物枝叶性状统计数据的影响。结果表明：（1）与传统的分类1相比,分类2对乔木植物枝叶性状影响的显著程度要高于分类3,而对灌木植物枝叶性状的影响程度低于分类3;但不同生长型分类方案中乔木和灌木的枝叶性状总体差异不显著。而与分类2小乔木相比,分类1以及分类2内部的乔木和灌木生长型的性状与分类2小乔木差异非常明显;（2）将不同生长型植物再划分为不同生活型后,不同生长型分类方案对性状统计的影响增大。无论是常绿还是落叶生活型的小乔木,其与不同生活型乔木和灌木的性状差异仍然显著。可见,不同的生长型分类方案可造成植物功能性状统计的差异;把小乔木植物这一功能类群划分出来,能更好地反映森林生态系统性状特征的差异性。     

The genetical theory of multilevel selection

The theory of multilevel selection (MLS) is beset with conceptual difficulties. Although it is widely agreed that covariance between group trait and group fitness may arise in the natural world and drive a response to ‘group selection’, ambiguity exists over the precise meaning of group trait and group fitness and as to whether group selection should be defined according to changes in frequencies of different types of individual or different types of group. Moreover, the theory of MLS has failed to properly engage with the problem of class structure, which greatly limits its empirical application to, for example, social insects whose colonies are structured into separate age, sex, caste and ploidy classes. Here, I develop a genetical theory of MLS, to address these problems. I show that taking a genetical approach facilitates a decomposition of group‐level traits – including reproductive success – into the separate contributions made by each constituent individual, even in the context of so‐called emergence. However, I uncover a novel problem with the group‐oriented approach: in many scenarios, it may not be possible to express a meaningful covariance between trait and fitness at the level of the social group, because the group's constituents belong to separate, irreconcilable classes.