Evolutionary trends of reproductive phenotype in Cycadales: an analysis of morphological evolution in Ceratozamia.

BACKGROUND AND AIMS: The size and shape of reproductive structures is especially relevant in evolution because these characters are directly related to the capacity of pollination and seed dispersal, a process that plays a basic role in evolutionary patterns. The evolutionary trajectories of reproductive phenotypes in gymnosperms have received special attention in terms of pollination and innovations related to the emergence of the Spermatophytes. However, variability of reproductive structures, evolutionary trends and the role of environment in the evolution of cycad species have not been well documented and explored. This study considered this topic under an explicitly phylogenetic and evolutionary approach that included a broad sampling of reproductive structures in the genus Ceratozamia. METHODS: We sampled 1400 individuals of 36 Ceratozamia species to explore evolutionary pattern and identify and evaluate factors that potentially drove their evolution. We analyzed characters for both pollen and ovulate strobili within a phylogenetic framework using different methods and characters (i. e., molecular and both quantitative and qualitative morphological) to infer phylogenetic relationships. Using this phylogenetic framework, evolutionary models of trait evolution for strobilar size were evaluated. In addition, quantitative morphological variation and its relation to environmental variables across species were analyzed. KEY RESULTS: We found contrasting phylogenetic signals between characters of pollen and ovulate strobili. These structures exhibited high morphological disparity in several characters related to size. Results of analyses of evolutionary trajectories suggested a stabilizing selection model. In regards to phenotype-environment, the analysis produced mixed results and differences for groups in the vegetation type where the species occur; however, a positive relationship with climatic variables was found. CONCLUSIONS: The integrated approach synthesized reproductive phenotypic variation with current phylogenetic hypotheses and provided explicit statements of character evolution. The characters of volume for ovulate strobili were the most informative, which could provide a reference for further study of the evolutionary complexity in Ceratozamia. Finally, heterogeneous environments, which are under changing weather conditions, promote variability of reproductive structures.

Testing the Tropical Niche Conservatism Hypothesis: Climatic Niche Evolution of Escallonia Mutis ex L. F. (Escalloniaceae).

We assess the Tropical Niche Conservatism Hypothesis in the genus Escallonia in South America using phylogeny, paleoclimate estimation and current niche modelling. We tested four predictions: (1) the climatic condition where the ancestor of Escallonia grew is megathermal; (2) the temperate niche is a derived condition from tropical clades; (3) the most closely related species have a similar current climate niche (conservation of the phylogenetic niche); and (4) there is a range expansion from the northern Andes to high latitudes during warm times. Our phylogenetic hypothesis shows that Escallonia originated 52.17 ± 0.85 My, in the early Eocene, with an annual mean temperature of 13.8 °C and annual precipitation of 1081 mm, corresponding to a microthermal to mesothermal climate; the species of the northern and central tropical Andes would be the ancestral ones, and the temperate species evolved between 32 and 20 My in a microthermal climate. The predominant evolutionary models were Brownian and Ornstein-Uhlenbeck. There was phylogenetic signal in 7 of the 9 variables, indicating conservation of the climatic niche. Escallonia would have originated in the central and southern Andes and reached the other environments by dispersion.

Reproductive and environmental traits explain the variation in egg size among Medusozoa (Cnidaria).

Medusozoa (Cnidaria) are characterized by diverse life cycles, with different semaphoronts (medusa, medusoid, fixed gonophore, polyp) representing the sexual phase and carrying the gametes. Although egg size is often considered a proxy to understand reproductive and developmental traits of medusozoans, understanding of the processes influencing egg size variation in the group under an evolutionary context is still limited. We carried out a comprehensive review of the variation of egg size in Medusozoa to test whether this variation is related to biological/sexual or environmental traits. Egg size presents a strong phylogenetic signal (λ = 0.79, K = 0.67), explaining why closely related species with different reproductive strategies and different individual sizes have similar egg sizes. However, variation in egg size is influenced by the number of eggs, depth and temperature, with larger eggs frequently present in species with few eggs (1-15), in deep-sea species and in cold-water species. Conversely, the production of small eggs among cold-water species of Staurozoa might be associated with the development of a small benthic larvae in this group. Our study reinforces that egg sizes respond to reproductive and environmental traits, although egg size is highly conserved within medusa classes.

Evolution of territoriality in Hylinae treefrogs: Ecological and morphological correlates and lineage diversification.

Given the diverse nature of traits involved in territorial defence, they may respond to different selective pressures and then exhibit distinct patterns of evolution. These selective pressures also may cause territorial behaviour to be associated with environmental and morphological variables. Such associations, however, have mostly been studied at the intraspecific level, being phylogenetic analyses of territoriality in a broad taxonomic framework rare in the literature. We used the anuran subfamily Hylinae to test (1) whether two territorial-behaviour traits with different levels of aggression-territorial call and physical combat-are evolutionarily more labile than a morphological trait used in physical combat-the spine-shaped prepollex; (2) whether reproduction in lentic waters and phytotelmata, as well as resource scarcity, might favour the occurrence of territoriality; (3) if physical combat is more important than territorial call for the evolution of body size and sexual size dimorphism and (4) the relationships between territorial-behaviour traits and lineage diversification. We mainly used the literature to build two datasets with different levels of certainty. Territorial-behaviour traits exhibited intermediate levels of phylogenetic signal in Hylinae, whereas the phylogenetic signal for the presence of the spine-shaped prepollex was strong. We found support for the hypothesis that reproduction in lentic water favours the occurrence of territorial behaviour, because the expression of territorial-behaviour traits was more associated with reproduction in lentic than in lotic waters. Territorial-behaviour traits were not correlated with annual precipitation nor with habitat complexity. Body size and sexual size dimorphism were not correlated with the presence of territorial call nor with physical combat. We identified negative correlations between diversification rates and physical combat. Relationships of territorial call and physical combat with diversification rates suggest that these territorial behaviours influence evolutionary processes in different ways.

Patterns of genome size evolution versus fraction of repetitive elements in statu nascendi species: the case of the willistoni subgroup of Drosophila (Diptera, Drosophilidae).

Genome size evolution is known to be related with transposable elements, yet such relation in incipient species remains poorly understood. For decades, the willistoni subgroup of Drosophila has been a model for evolutionary studies because of the different evolutionary stages and degrees of reproductive isolation its species present. Our main question here was how speciation influences genome size evolution and the fraction of repetitive elements, with a focus on transposable elements. We quantitatively compared the mobilome of four species and two subspecies belonging to this subgroup with their genome size, and performed comparative phylogenetic analyses. Our results showed that genome size and the fraction of repetitive elements evolved according to the evolutionary history of these species, but the content of transposable elements showed some discrepancies. Signals of recent transposition events were detected for different superfamilies. Their low genomic GC content suggests that in these species transposable element mobilization might be facilitated by relaxed natural selection. Additionally, a possible role of the superfamily DNA/TcMar-Tigger in the expansion of these genomes was also detected. We hypothesize that the undergoing process of speciation could be promoting the observed increase in the fraction of repetitive elements and, consequently, genome size.

Investigating the diversification of holocentromeric satellite DNA Tyba in Rhynchospora (Cyperaceae).

BACKGROUND AND AIMS: Satellite DNAs (satDNAs) are repetitive sequences composed by tandemly arranged, often highly homogenized units called monomers. Although satDNAs are usually fast evolving, some satDNA families can be conserved across species separated by several millions of years, probably because of their functional roles in the genomes. Tyba was the first centromere-specific satDNA described for a holocentric organism, until now being characterized for only eight species of the genus Rhynchospora Vahl. (Cyperaceae). Here, we characterized Tyba across a broad sampling of the genus, analysing and comparing its evolutionary patterns with other satDNAs. METHODS: We characterized the structure and sequence evolution of satDNAs across a robust dadated phylogeny based on Hybrid Target-Capture Sequencing (hyb-seq) of 70 species. We mined the repetitive fraction for Tyba-like satellites to compare its features with other satDNAs and to construct a Tyba-based phylogeny for the genus. KEY RESULTS: Our results show that Tyba is present in the majority of examined species of the genus, spanning four of the five major clades and maintaining intrafamily pairwise identity of 70.9% over 31 Myr. In comparison, other satellite families presented higher intrafamily pairwise identity but are phylogenetically restricted. Furthermore, Tyba sequences could be divided into 12 variants grouped into three different clade-specific subfamilies, showing evidence of traditional models of satDNA evolution, such as the concerted evolution and library models. Besides, a Tyba-based phylogeny showed high congruence with the hyb-seq topology. Our results show structural indications of a possible relationship of Tyba with nucleosomes, given its high curvature peaks over conserved regions and overall high bendability values compared with other non-centromeric satellites. CONCLUSIONS: Overall, Tyba shows a remarkable sequence conservation and phylogenetic significance across the genus Rhynchospora, which suggests that functional roles might lead to long-term stability and conservation for satDNAs in the genome.

Annual precipitation predicts the phylogenetic signal in bat-fruit interaction networks across the Neotropics.

Closely related species tend to be more similar than randomly selected species from the same phylogenetic tree. This pattern, known as a phylogenetic signal, has been extensively studied for intrinsic (e.g. morphology), as well as extrinsic (e.g. climatic preferences), properties but less so for ecological interactions. Phylogenetic signals of species interactions (i.e. resource use) can vary across time and space, but the causes behind such variations across broader spatial extents remain elusive. Here, we evaluated how current and historical climates influence phylogenetic signals of bat-fruit interaction networks across the Neotropics. We performed a model selection relating the phylogenetic signals of each trophic level (bats and plants) with a set of current and historical climatic factors deemed ecologically important in shaping biotic interactions. Bat and plant phylogenetic signals in bat-fruit interaction networks varied little with climatic factors, although bat phylogenetic signals positively covaried with annual precipitation. These findings indicated that water availability could increase resource availability, favouring higher niche partitioning of trophic resources among bat species and hence bat phylogenetic signals across bat-fruit interaction networks. Overall, our study advances our understanding of the spatial dynamics of bat-fruit interactions by highlighting the association of current climatic factors with phylogenetic patterns of biotic interactions.

The evolution of red blood cell shape in fishes.

The size and shape of red blood cells (RBCs) provide key information on life-history strategies in vertebrates. However, little is known about how RBC shape evolved in response to environmental factors, body size and the role of evolutionary rate. Here, we analysed RBC morphometrics in a set of Teleostei (bony fishes) and Elasmobranchii (sharks and rays) species testing the hypothesis that phylogenetic relationship explains species occupation of morphospace. We collected data on cell and nucleus area and volume, nucleus:cytoplasm ratio and shape factor for 65 species belonging to 28 orders. Then, we built phylomorphospaces separately for bony fish and sharks and rays. To test whether phylogenetic relationships predicted phenotypic similarity, we calculated multivariate phylogenetic signal. We also estimated the evolutionary rate of RBC shape for each node and tip using ridge regression. Finally, we tested whether habitat and body size influenced RBC shape using a PGLS. We found a significant phylogenetic signal in RBC shape for bony fish, but not sharks and rays. Saltwater teleost species were more clustered than freshwater ones in the phylomorphospace, suggesting clade disparity. Accordingly, the rate of evolution was highly heterogeneous, with significant decrease in Acanthopterygii. Neither habitat nor body size influenced RBC shape. In conclusion, RBC shape seems to have evolved in fishes in response to multiple selective pressures independent of life-history characters.

Timber and non-timber forest products in the northernmost Neotropical rainforest: Ecological factors unravel their landscape distribution.

The benefits provided by tropical rainforests are unevenly distributed throughout the landscape and are shaped by abiotic and biotic components that influence the spatial distribution and functional traits of the species involved. We tested whether environmental stratification of the rainforest in biophysical Landscape Units (LU), defined by topography and soil, is related to the spatial distribution of diversity, abundance and productivity (standing biomass) of tree assemblages that provide potential forest products (PFP). Considering that different PFP are associated with specific plant traits, we also tested whether a phylogenetic signal exists among the species that comprise specific use categories. Non-metric multidimensional scaling ordinations and permutational analysis of variance were based on the frequency, abundance and productivity of 129 species, the PFP of which were classified as fodder, food, fuelwood, medicinal, melliferous, ornamental, plywood and timber in 15 plots of 0.5 ha each. We constructed a phylogenetic tree of the studied species and analyzed the phylogenetic signal strength (D-statistic) among them. The spatial distribution of diversity and abundance of useful species changes among the LU. Specific PFP can be provided in contrasting habitat conditions, but generally not by the same species. The PFP categories that presented a phylogenetic signal were associated with wood characteristics (fuelwood and plywood) and the palatability of the leaves and reproductive structures (fodder). The Moraceae family was significantly related to fodder and plywood, whereas Meliaceae, Myrtaceae and Sapotaceae were mostly used for fuelwood. The medicinal species presented convergent traits distributed throughout the phylogeny. However, since our study included a broad variety of plant structures, it is possible that phylogenetic dispersion can change if we consider the specific uses within each category. Our findings show that the assemblages of PFP suppliers can be clustered through biophysical units based on soil and topography, and specific categories of PFP are often supplied by phylogenetically related species. This knowledge is fundamental in order to incorporate the high diversity of tree species and their potential uses into productive reforestation and agroforestry programs.

Elevation does not strongly influence interspecific variation in body size of small Tropical endotherms / La elevación no influencia fuertemente la variación interespecífica en el tamaño corporal de los endotermos tropicales pequeños

Introduction: Body size is an essential trait for endotherms to face the physiological requirements of cold, so there is a tendency to large body size at high altitudes and latitudes, known as Bergmann's rule. However, the validity of this ecomorphological rule to small-bodied endotherms across altitudinal gradients is poorly known. Objective: To understand the effects of environmental variation on body size, we assessed whether interspecific variation in body size of small tropical endotherms follows Bergmann's rule along tropical altitudinal gradients. Methods: We compiled data on elevational ranges and body masses for 133 species of hummingbirds of Colombia. We then assessed the association between body mass and mid-point of the altitudinal distribution using phylogenetic generalized least squares (PGLS) analyses under different evolutionary models. Results: We found a decelerating rate of evolution for body size since the Early Burst model of evolution provided a better fit to body mass data. For elevational range, we found a slow and constant rate since Pagel's lambda model provided a better fit to the mid-point of the altitudinal distribution data. Besides, phylogenetic regression analysis indicated that body mass and the altitudinal range of hummingbirds are associated through the phylogeny, with a positive but slight association (R2= 0.036). Conclusions: We found that body mass and altitude of hummingbirds are positively related, which is in agreement with expectations under Bergmann's rule. However, this association was weaker than expected for small and non-passerine birds like hummingbirds. Thus, our results suggest that environmental changes across altitudinal gradients do not strongly influence body mass in small tropical endotherms as hummingbirds.

Introducción: El tamaño corporal es un rasgo importante para determinar la respuesta de los endotermos a los requerimientos que exigen las zonas frías, por lo cual se espera una tendencia hacia el incremento del tamaño corporal al aumentar la altitud y la latitud. Sin embargo, se conoce poco acerca de la validez de esta regla ecomorfológica, conocida como la regla de Bergmann, para endotermos pequeños en gradientes altitudinales tropicales. Objetivo: Con el fin de entender los efectos de la variación ambiental sobre el tamaño corporal, se evaluó sí la variación interespecífica en la masa corporal de endotermos tropicales pequeños se ajusta a la regla de Bergmann a lo largo de gradientes de elevación. Métodos: Se compilaron datos sobre los rangos de distribución altitudinal y los tamaños corporales de 133 especies de colibríes en Colombia. Posteriormente, se evaluó la asociación entre la masa corporal y el punto medio de distribución altitudinal de los colibríes mediante análisis de mínimos cuadrados generalizados filogenéticos (PGLS) bajo diferentes modelos evolutivos. Resultados: La evolución de la masa corporal se ajustó mejor a un modelo de evolución Early Burst, mientras que el rango de elevación al modelo evolutivo lambda de Pagel; lo que indica que la tasa de evolución es desacelerada para el tamaño del cuerpo, mientras es lenta y constante para el rango de elevación. Además, el análisis de regresión filogenética indica que la masa corporal y el rango de elevación están positiva y ligeramente asociados (R2 = 0.036). Conclusiones: De acuerdo con lo esperado por la regla de Bergmann, los resultados indican que los colibríes tienden a ser más grandes a mayores altitudes. Sin embargo, esta asociación es más débil de lo esperado para aves no paseriformes de tamaño pequeño como los colibríes.Por lo tanto, los resultados sugieren que las variaciones ambientales a lo largo de gradientes de elevación no tienen una influencia fuerte sobre el tamaño corporal de endotermos pequeños como los colibríes.

The use of wing shape for characterising macroevolution in mosquitoes (Diptera: Culicidae)

The wing form of culicid mosquitoes shows considerable variation among groups: this phenomenon has been addressed by several studies through space-time analyses in mosquito populations, species, and genera. The observed variation results from a combination of two distinct factors: heredity and phenotypic plasticity. The first is usually related to wing shape, a complex character that may serve as a taxonomic marker in specific cases. We hypothesized that wing shape might be phylogenetically meaningful in Culicidae. In this study, we applied a geometric morphometrical approach based on 18 landmarks in 81 species of mosquitoes, representing 19 different genera, to investigate whether wing shape can help retrieve macroevolutionary patterns or identify any phylogenetic signals. We observed that wing shape differed considerably among groups, especially between Anophelinae and Culicinae subfamilies; thus, some wing shape elements may be synapomorphic. Comparisons among wing consensus after Procrustes superimposition revealed that landmark #1, located between the veins RS and R1, was the most variable. Sabethini tribe was distinguished from other taxa owing to a strong phylogenetic signal of its wings, whereas other culicids presented weaker signals and were not that distinguishable. Evolutionary forces such as natural selection, evolutionary limitation/constraint, or canalization mechanisms might drive the evolution of wing phenotype. These findings suggest that the wing undergoes evolution over long periods, but is not neutral enough to reconstruct the phylogenetic history of these insects. Gene-based studies should be performed to understand the driving forces in wing evolution.

Megafauna Seed Dispersal in the Neotropics: A Meta-Analysis Shows No Genetic Signal of Loss of Long-Distance Seed Dispersal.

Restricted gene flow may lead to the loss of genetic diversity and higher genetic differentiation among populations, but the genetic consequences of megafauna extinction for plant populations still remain to be assessed. We performed a phylogenetic-independent meta-analysis across 102 Neotropical plants to test the hypothesis that plant species with megafaunal seed dispersal syndrome have a lower genetic diversity and a higher genetic differentiation than those without it. We classified as megafauna-dependent plant species those that potentially relied only on megafauna to seed dispersal, and as megafauna-independent those that relied on megafauna and other seed dispersers. Our data comprised 98 studies using microsatellite markers. We found no statistical difference in genetic diversity and differentiation between plants with megafauna and non-megafauna seed dispersal syndrome, although the statistical power to detect differences in genetic differentiation was low. Moreover, we found no statistical difference between megafauna-dependent and megafauna-independent plant species. We then used generalized linear mixed models and phylogenetic generalized least square models to investigate the effects of megafaunal seed dispersal syndromes and reproductive traits on variation in genetic diversity and genetic differentiation. We found no effect of megafaunal syndrome, rather, reproductive traits, such as pollination mode, mating, and breeding systems, showed significant effects. Our findings show that the genetic studies of Neotropical plants performed so far show no difference in genetic diversity and differentiation in plants with megafaunal compared to those with non-megafaunal seed dispersal syndromes. Our results also provide evidence pointing out that plant species with megafaunal seed dispersal syndromes may have used different strategies to counterbalance the extinction of their mutualistic megafauna dispersers, such as the dispersal by extant mammals that may promote long-distance seed dispersal. Our results also reinforce the importance of pollination to long-distance gene flow in Neotropical plants.

Evolution of Ceratozamia cycads: A proximate-ultimate approach.

Evidence suggests that past climatic fluctuations affected speciation of extant cycads. However, empirical genetic and morphological evidence explaining patterns and processes of species diversification are scarce. There are some explanations for the origin and evolution of the genus Ceratozamia, but with inconclusive results. To elucidate the evolution of Ceratozamia, we used genetic and phenotypic sources as empirical data, which were applied in a 'proximate-ultimate' framework (ecological and evolutionary scale, respectively). Our results suggested that the evolutionary mechanisms of speciation were shaped by deterministic (natural selection-adaptation) driven by climatic conditions associated to water stress, and probably enhanced by stochastic processes (gene drift and inbreeding). In general terms, punctuated evolution models were those that best explained the patterns of speciation throughout the phylogenetic history of the lineages encompassed in the genus Ceratozamia. Finally, we provide empirical evidence on the tempo and mode of the evolution of a 'living plant fossil'.

Phylogenetic signal analysis in the basicranium of Ursidae (Carnivora, Mammalia).

Ursidae is a monophyletic group comprised of three subfamilies: Tremarctinae, Ursinae and Ailuropodinae, all of which have a rich geographical distribution. The phylogenetic relationships within the Ursidae group have been underexamined, especially regarding morphological traits such as the basicranium. Importantly, the basicranium is a highly complex region that covers a small portion of the skull, combining both structural and functional aspects that determine its morphology. Phylogenetic hypotheses of the Ursidae (including Tremarctinae) have been made based on morphological characters that considers skull, mandible and teeth features, while specific characters of the auditory region and basicranium have not been taken into account. To do this, we analyse the shape and size macroevolution of the basicranium of Ursidae, testing its morphological disparity in a phylogenetic context, which is quantified by means of the phylogenetic signal. We investigated phylogenetical autocorrelation by shape (depicted by Principal Components Analysis scores from previous published analyses) and basicranium size (depicted by centroid size, CS) using an orthonormal decomposition analysis and Abouheif C mean. The main advantages of these methods are that they rely exclusively on cladogram topology and do not require branch-length estimates. Also, an optimisation of the ancestral nodes was performed using TNT 1.5 software. In relation to the phylogenetic signal, both methods showed similar results: the presence of autocorrelation was detected in PC1 and PC2, while in PC3, PC4 and PC5 and in the size of the basicranium (CS), the absence of autocorrelation occurred. The most significant nodes (where there is autocorrelation) are the basal nodes 'Ursidae' and 'Ursinae-Tremarctinae'. Within this last group, distinctive basicranium morphology is observed, being more conservative in Tremarctinae than in Ursinae. The differences between these subfamilies could be related to historical events involving varying food and environmental preferences. The high phylogenetic signal in the node Tremarctinae probably indicates that the basicranium configuration of these bears was obtained early in their evolutionary history. Finally, our results of the basicranium and skull length ratios indicate that in Tremarctinae, the basicranium size was not determined by phylogeny but instead by other factors, such as adaptive responses to climatic changes and competition with other carnivores.

Looking beyond the flowers: associations of stingless bees with sap-sucking insects.

The main sources of food for stingless bees are the nectar and pollen harvested from flowers, whereas one important kind of nesting material (i.e. wax) is produced by their own abdominal glands. Stingless bees can, nonetheless, obtain alternative resources of food and wax from exudates released by sap-sucking insects as honeydew and waxy cover, respectively. To date, there are no comprehensive studies investigating how diversified and structured the network interactions between stingless bees and sap-sucking insects are. Here, we conducted a survey of the data on relationship between stingless bees and sap-sucking insects to evaluate: (1) which resources are collected by which stingless bee species; (2) how diverse the interaction network is, using species degree and specialisation index as a proxy; and if (3) there would be any phylogenetic signal in the species degree and specialisation indices. Our findings demonstrate that approximately 21 stingless bee species like Trigona spp. and Oxytrigona spp. have been observed interacting with 11 sap-sucking species, among which Aethalion reticulatum is the main partner. From ca. 50 records, Brazil is the country with most observations (n = 38) of this type of ecological interaction. We found also that stingless bees harvest fivefold more honeydew than waxy covers on sap-sucking insects. However, we did not find any phylogenetic signal for the occurrence of this interaction, considering species degree and specialisation indices, suggesting that both traits apparently evolved independently among stingless bee species. We suggest that specific ecological demands may drive this opportunistic behaviour exhibited by stingless bees, because major sources of food are obtained from flowers and these bees produce their own wax.

Phylogeny, fruit traits, and ecological correlates of fruiting phenology in a Neotropical dry forest.

In tropical dry forests, a high interspecific variation in the strategies of fruiting phenology has been documented. Therefore, phenological responses may be mediated by influence of environmental variables, functional plant attributes or phylogenetic inertia. During 2 years, we recorded the fruiting phenology of 151 species belonging to 5 different growth forms of a Neotropical dry forest in Mexico. We evaluated the relationships between fruiting phenology, abiotic factors (precipitation, temperature, day-length) and functional attributes (growth form, dispersal syndrome, size and time for fruit development) using phylogenetic least squares models (PGLS). More species had ripe fruits during the dry season (92%) than during rainy months and dispersed their seeds by autochory and endozoochory. We found that fruit development time was positively correlated with fruit size and together the morphological fruit traits (size and dispersal syndrome) showed an important relationship with the growth form, but with a strong phylogenetic signal. Environmental seasonality had a strong influence on fruit ripening time, without a relevant association to the phylogeny of plant species. However, the phenological response to the environment (rainfall and day-length) at the community level was mediated by growth form. In woody species, we documented a high interspecific fruiting variation linked with the different dispersal syndromes. In herbaceous species, fruiting phenology is a trait restricted by the duration of their life cycle by rainfall seasonality, which in turn might have selected some traits (e.g., dry fruit, presence of spines, explosive dehiscence) for maximizing seed dispersal during the dry season.

Evolution of Climatic Related Leaf Traits in the Family Nothofagaceae.

The current relationship between leaf traits and environmental variables has been widely used as a proxy for climate estimates. However, it has been observed that the phylogenetic relationships between taxa also influence the evolution of climatic related leaf traits, implying that the direct use of the physiognomy-climate relation should be corrected by their ancestor-descendant relations. Here, we analyze the variation of 20 leaf traits during the evolution of 27 species in the Gondwana family Nothofagaceae. We evaluate whether the evolution of these traits is exclusively associated with past climate variations or whether they are restricted by phylogenetic relationships. Our results indicate that four leaf traits, associated with size and shape, had consistently a phylogenetic independent evolution, suggesting adaptive variation with the environment. While three of the traits, presented consistently phylogenetic signal and fit a Brownian motion or Ornstein-Uhlenbeck model of evolution, suggesting that the evolution of these traits is restrained by phylogenetic relationships and implying that phylogenetic corrections should be made for the family Nothofagaceae to use them as climatic proxy. Finally, this study highlights the importance of evaluating the evolutionary history of climatic related leaf traits before conducting paleoclimate estimates.

Interspecific geographic range size-body size relationship and the diversification dynamics of Neotropical furnariid birds.

Among the earliest macroecological patterns documented, is the range and body size relationship, characterized by a minimum geographic range size imposed by the species' body size. This boundary for the geographic range size increases linearly with body size and has been proposed to have implications in lineages evolution and conservation. Nevertheless, the macroevolutionary processes involved in the origin of this boundary and its consequences on lineage diversification have been poorly explored. We evaluate the macroevolutionary consequences of the difference (hereafter the distance) between the observed and the minimum range sizes required by the species' body size, to untangle its role on the diversification of a Neotropical species-rich bird clade using trait-dependent diversification models. We show that speciation rate is a positive hump-shaped function of the distance to the lower boundary. The species with highest and lowest distances to minimum range size had lower speciation rates, while species close to medium distances values had the highest speciation rates. Further, our results suggest that the distance to the minimum range size is a macroevolutionary constraint that affects the diversification process responsible for the origin of this macroecological pattern in a more complex way than previously envisioned.

Analyzing community-weighted trait means across environmental gradients: should phylogeny stay or should it go?

Functional traits mediate ecological responses of organisms to the environment, determining community structure. Community-weighted trait means (CWM) are often used to characterize communities by combining information on species traits and distribution. Relating CWM variation to environmental gradients allows for evaluating species sorting across the metacommunity, either based on correlation tests or ordinary least squares (OLS) models. Yet, it is not clear if phylogenetic signal in both traits and species distribution affect those analyses. On one hand, phylogenetic signal might indicate niche conservatism along clade evolution, reinforcing the environmental signal in trait assembly patterns. On the other hand, it might introduce phylogenetic autocorrelation to mean trait variation among communities. Under this latter scenario, phylogenetic signal might inflate type I error in analysis relating CWM variation to environmental gradients. We explore multiple ways phylogenetic history may influence analysis relating CWM to environmental gradients. We propose the concept of neutral trait diffusion, which predicts that for a functional trait x, CWM variation among local communities does not deviate from the expectation that x evolved according to a neutral evolutionary process. Based on this framework we introduce a graphical tool called neutral trait diffusion representation (NTDR) that allows for the evaluation of whether it is necessary to carry out phylogenetic correction in the trait prior to analyzing the association between CWM and environmental gradients. We illustrate the NTDR approach using simulated traits, phylogenies and metacommunities. We show that even under moderate phylogenetic signal in both the trait used to define CWM and species distribution across communities, OLS models relating CWM variation to environmental gradients lead to inflated type I error when testing the null hypothesis of no association between CWM and environmental gradient. To overcome this issue, we propose a phylogenetic correction for OLS models and evaluate its statistical performance (type I error and power). Phylogeny-corrected OLS models successfully control for type I error in analysis relating CWM variation to environmental gradients but may show decreased power. Combining the exploratory tool of NTDR and phylogenetic correction in traits, when necessary, guarantees more precise inferences about the environmental forces driving trait-mediated species sorting across metacommunities.

Coevolution between flight morphology, vertical stratification and sexual dimorphism: what can we learn from tropical butterflies?

Occurrence patterns are partly shaped by the affinity of species with habitat conditions. For winged organisms, flight-related attributes are vital for ecological performance. However, due to the different reproductive roles of each sex, we expect divergence in flight energy budget, and consequently different selection responses between sexes. We used tropical frugivorous butterflies as models to investigate coevolution between flight morphology, sex dimorphism and vertical stratification. We studied 94 species of Amazonian fruit-feeding butterflies sampled in seven sites across 3341 ha. We used wing-thorax ratio as a proxy for flight capacity and hierarchical Bayesian modelling to estimate stratum preference. We detected a strong phylogenetic signal in wing-thorax ratio in both sexes. Stouter fast-flying species preferred the canopy, whereas more slender slow-flying species preferred the understorey. However, this relationship was stronger in females than in males, suggesting that female phenotype associates more intimately with habitat conditions. Within species, males were stouter than females and sexual dimorphism was sharper in understorey species. Because trait-habitat relationships were independent from phylogeny, the matching between flight morphology and stratum preference is more likely to reflect adaptive radiation than shared ancestry. This study sheds light on the impact of flight and sexual dimorphism on the evolution and ecological adaptation of flying organisms.