Evolutionary transitions in diet influence the exceptional diversification of a lizard adaptive radiation.

BACKGROUND: Diet is a key component of a species ecological niche and plays critical roles in guiding the trajectories of evolutionary change. Previous studies suggest that dietary evolution can influence the rates and patterns of species diversification, with omnivorous (animal and plant, 'generalist') diets slowing down diversification compared to more restricted ('specialist') herbivorous and carnivorous diets. This hypothesis, here termed the "dietary macroevolutionary sink" hypothesis (DMS), predicts that transitions to omnivorous diets occur at higher rates than into any specialist diet, and omnivores are expected to have the lowest diversification rates, causing an evolutionary sink into a single type of diet. However, evidence for the DMS hypothesis remains conflicting. Here, we present the first test of the DMS hypothesis in a lineage of ectothermic tetrapods-the prolific Liolaemidae lizard radiation from South America. RESULTS: Ancestral reconstructions suggest that the stem ancestor was probably insectivorous. The best supported trait model is a diet-dependent speciation rate, with independent extinction rates. Herbivory has the highest net diversification rate, omnivory ranks second, and insectivory has the lowest. The extinction rate is the same for all three diet types and is much lower than the speciation rates. The highest transition rate was from omnivory to insectivory, and the lowest transition rates were between insectivory and herbivory. CONCLUSIONS: Our findings challenge the core prediction of the DMS hypothesis that generalist diets represent an 'evolutionary sink'. Interestingly, liolaemid lizards have rapidly and successfully proliferated across some of the world's coldest climates (at high elevations and latitudes), where species have evolved mixed arthropod-plant (omnivore) or predominantly herbivore diets. This longstanding observation is consistent with the higher net diversification rates found in both herbivory and omnivory. Collectively, just like the evolution of viviparity has been regarded as a 'key adaptation' during the liolaemid radiation across cold climates, our findings suggest that transitions from insectivory to herbivory (bridged by omnivory) are likely to have played a role as an additional key adaptation underlying the exceptional diversification of these reptiles across extreme climates.

Disturbance reinforces community assembly processes differentially across spatial scales.

BACKGROUND AND AIMS: There is a paucity of empirical research and a lack of predictive models concerning the interplay between spatial scale and disturbance as they affect the structure and assembly of plant communities. We proposed and tested a trait dispersion-based conceptual model hypothesizing that disturbance reinforces assembly processes differentially across spatial scales. Disturbance would reinforce functional divergence at the small scale (neighbourhood), would not affect functional dispersion at the intermediate scale (patch) and would reinforce functional convergence at the large scale (site). We also evaluated functional and species richness of native and exotic plants to infer underlying processes. Native and exotic species richness were expected to increase and decrease with disturbance, respectively, at the neighbourhood scale, and to show similar associations with disturbance at the patch (concave) and site (negative) scales. METHODS: In an arid shrubland, we estimated species richness and functional dispersion and richness within 1 m2 quadrats (neighbourhood) nested within 100 m2 plots (patch) along a small-scale natural disturbance gradient caused by an endemic fossorial rodent. Data for the site scale (2500 m2 plots) were taken from a previous study. We also tested the conceptual model through a quantitative literature review and a meta-analysis. KEY RESULTS: As spatial scale increased, disturbance sequentially promoted functional divergence, random trait dispersion and functional convergence. Functional richness was unaffected by disturbance across spatial scales. Disturbance favoured natives over exotics at the neighbourhood scale, while both decreased under high disturbance at the patch and site scales. CONCLUSIONS: The results supported the hypothesis that disturbance reinforces assembly processes differentially across scales and hampers plant invasion. The quantitative literature review and the meta-analysis supported most of the model predictions.

The importance of scale-dependent ravine characteristics on breeding-site selection by the Burrowing Parrot, Cyanoliseus patagonus.

In birds, the environmental variables and intrinsic characteristics of the nest have important fitness consequences through its influence on the selection of nesting sites. However, the extent to which these variables interact with variables that operate at the landscape scale, and whether there is a hierarchy among the different scales that influences nest-site selection, is unknown. This interaction could be crucial in burrowing birds, which depend heavily on the availability of suitable nesting locations. One representative of this group is the burrowing parrot, Cyanoliseus patagonus that breeds on specific ravines and forms large breeding colonies. At a particular site, breeding aggregations require the concentration of adequate environmental elements for cavity nesting, which are provided by within ravine characteristics. Therefore, intrinsic ravine characteristics should be more important in determining nest site selection compared to landscape level characteristics. Here, we assess this hypothesis by comparing the importance of ravine characteristics operating at different scales on nest-site selection and their interrelation with reproductive success. We quantified 12 characteristics of 105 ravines in their reproductive habitat. For each ravine we quantified morphological variables, distance to resources and disturbance as well as nest number and egg production in order to compare selected and non-selected ravines and determine the interrelationship among variables in explaining ravine differences. In addition, the number of nests and egg production for each reproductive ravine was related to ravine characteristics to assess their relation to reproductive success. We found significant differences between non-reproductive and reproductive ravines in both intrinsic and extrinsic characteristics. The multidimensional environmental gradient of variation between ravines, however, shows that differences are mainly related to intrinsic morphological characteristics followed by extrinsic variables associated to human disturbance. Likewise, within reproductive ravines, intrinsic characteristics are more strongly related to the number of nests. The probability of producing eggs, however, was related only to distance to roads and human settlements. Patterns suggest that C. patagonus mainly selects nesting sites based on intrinsic morphological characteristics of ravines. Scale differences in the importance of ravine characteristics could be a consequence of the particular orography of the breeding habitat. The arrangement of resources is associated to the location of the gullies rather than to individual ravines, determining the spatial availability and disposition of resources and disturbances. Thus, nest selection is influenced by intrinsic characteristics that maximize the fitness of individuals. Scaling in nest-selection is discussed under an optimality approach that partitions patch selection based on foraging theory.

Disturbance by an endemic rodent in an arid shrubland is a habitat filter: effects on plant invasion and taxonomical, functional and phylogenetic community structure.

Background and Aims: Disturbance often drives plant invasion and may modify community assembly. However, little is known about how these modifications of community patterns occur in terms of taxonomic, functional and phylogenetic structure. This study evaluated in an arid shrubland the influence of disturbance by an endemic rodent on community functional divergence and phylogenetic structure as well as on plant invasion. It was expected that disturbance would operate as a habitat filter favouring exotic species with short life cycles. Methods: Sixteen plots were sampled along a disturbance gradient caused by the endemic fossorial rodent Spalacopus cyanus , measuring community parameters and estimating functional divergence for life history traits (functional dispersion index) and the relative contribution to functional divergence of exotic and native species. The phylogenetic signal (Pagel's lambda) and phylogenetic community structure (mean phylogenetic distance and mean nearest taxon phylogenetic distance) were also estimated. The use of a continuous approach to the disturbance gradient allowed the identification of non-linear relationships between disturbance and community parameters. Key Results: The relationship between disturbance and both species richness and abundance was positive for exotic species and negative for native species. Disturbance modified community composition, and exotic species were associated with more disturbed sites. Disturbance increased trait convergence, which resulted in phylogenetic clustering because traits showed a significant phylogenetic signal. The relative contribution of exotic species to functional divergence increased, while that of natives decreased, with disturbance. Exotic and native species were not phylogenetically distinct. Conclusions: Disturbance by rodents in this arid shrubland constitutes a habitat filter over phylogeny-dependent life history traits, leading to phylogenetic clustering, and drives invasion by favouring species with short life cycles. Results can be explained by high phenotypic and phylogenetic resemblance between exotic and native species. The use of continuous gradients when studying the effects of disturbance on community assembly is advocated.

Woody climbers show greater population genetic differentiation than trees: Insights into the link between ecological traits and diversification.

The climbing habit is a key innovation in plants: climbing taxa have higher species richness than nonclimbing sister groups. We evaluated the hypothesis that climbing plant species show greater among-population genetic differentiation than nonclimber species. We compared the among-population genetic distance in woody climbers (eight species, 30 populations) and trees (seven species, 29 populations) coexisting in nine communities in a temperate rainforest. We also compared within-population genetic diversity in co-occurring woody climbers and trees in two communities. Mean genetic distance between populations of climbers was twice that of trees. Isolation by distance (increase in genetic distance with geographic distance) was greater for climbers. Climbers and trees showed similar within-population genetic diversity. Our longevity estimate suggested that climbers had shorter generation times, while other biological features often associated with diversification (dispersal and pollination syndromes, mating system, size, and metabolic rate) did not show significant differences between groups. We hypothesize that the greater population differentiation in climbers could result from greater evolutionary responses to local selection acting on initially higher within-population genetic diversity, which could be driven by neutral processes associated with shorter generation times. Increased population genetic differentiation could be incorporated as another line of evidence when testing for key innovations.

Species divergence and phylogenetic variation of ecophysiological traits in lianas and trees.

The climbing habit is an evolutionary key innovation in plants because it is associated with enhanced clade diversification. We tested whether patterns of species divergence and variation of three ecophysiological traits that are fundamental for plant adaptation to light environments (maximum photosynthetic rate [A(max)], dark respiration rate [R(d)], and specific leaf area [SLA]) are consistent with this key innovation. Using data reported from four tropical forests and three temperate forests, we compared phylogenetic distance among species as well as the evolutionary rate, phylogenetic distance and phylogenetic signal of those traits in lianas and trees. Estimates of evolutionary rates showed that R(d) evolved faster in lianas, while SLA evolved faster in trees. The mean phylogenetic distance was 1.2 times greater among liana species than among tree species. Likewise, estimates of phylogenetic distance indicated that lianas were less related than by chance alone (phylogenetic evenness across 63 species), and trees were more related than expected by chance (phylogenetic clustering across 71 species). Lianas showed evenness for R(d), while trees showed phylogenetic clustering for this trait. In contrast, for SLA, lianas exhibited phylogenetic clustering and trees showed phylogenetic evenness. Lianas and trees showed patterns of ecophysiological trait variation among species that were independent of phylogenetic relatedness. We found support for the expected pattern of greater species divergence in lianas, but did not find consistent patterns regarding ecophysiological trait evolution and divergence. R(d) followed the species-level pattern, i.e., greater divergence/evolution in lianas compared to trees, while the opposite occurred for SLA and no pattern was detected for A(max). R(d) may have driven lianas' divergence across forest environments, and might contribute to diversification in climber clades.

Co-occurrence patterns along a regional aridity gradient of the subtropical Andes do not support stress gradient hypotheses.

The stress gradient hypothesis posits that facilitation and stress are positively correlated. The hump-shaped hypothesis, on the contrary, proposes that facilitation is greater at intermediate stress levels. The relationship between facilitation and environmental stress is commonly studied at small spatial scales and/or considering few species; thus, the implications of facilitation at a community level remain poorly understood. Here, we analyzed local co-occurrence patterns of all plant species at 25 sites within the subtropical Andes to evaluate the role of facilitation and competition as drivers of community structure. We considered a wide latitudinal gradient (19-26°S) that incorporates great variation in aridity. No previous studies have attempted to study these patterns across such a broad scale in warm deserts. Each locality was sampled at two scales (quadrat and patch), and co-occurrence was analyzed via null models. Furthermore, we tested for a relationship between plant co-occurrences and environmental aridity. Resulting patterns depended on life form. When all species were considered, negative associations were found, indicating competition. Woody/cactus life forms tended to be associated across communities, suggesting that there is facilitation between these life forms. Additionally, and unlike previous studies, we found positive associations among shrubs. The strength of the association between woody species changed non-monotonically with aridity. Herbs showed an inverted hump-shaped relationship, albeit ranging mostly among neutral values. Independent of the association type exhibited by different life forms, our community level results do not support current stress gradient hypotheses.

Context-dependent recruitment of Guettarda viburnoides in a Neotropical forest-savanna mosaic.

PREMISE OF THE STUDY: Seed dispersal is a key process for plant regeneration in tropical areas. Differences in disperser behavior result in a nonrandom distribution of seeds among habitats. Patterns of seed dispersal may cascade through the entire recruitment phase or uncoupling between developmental stages may occur such that patterns of seed dispersal are discordant with patterns of recruitment. Here, we analyzed how habitat and interannual variability affect the recruitment dynamics of a bird-dispersed tree, Guettarda viburnoides (RUB.). METHODS: Over 3 years, we determined the habitats where seeds of G. viburnoides are dispersed, and we experimentally quantified postdispersal seed predation, seedling emergence, and survival in four habitats of a forest-savanna mosaic in Bolivia. KEY RESULTS: Habitat affected seed dispersal, seed predation, and seedling emergence. The strength of postdispersal processes, however, varied between years and no consistent within-habitat pattern emerged. Uncoupling among different life-stages was observed across habitats, and spatial concordance was found between seed rain and sapling recruitment patterns. CONCLUSIONS: Habitat can affect seed dispersal, postdispersal processes and the recruitment dynamics of a Neotropical tree in a heterogeneous landscape. Additionally, our results show interannual variability in the strength of postdispersal processes, which leads to shifts in habitat suitability between years. Therefore, to better understand the role of site suitability for recruitment, we need to explicitly consider not only variation in habitat-specificity for dispersal and postdispersal processes, but also how this variation can shift under different environmental conditions-that is, the context dependence of suitability.

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae).

The benefits of ant-plant-herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant-plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations.