How flower colour signals allure bees and hummingbirds: a community-level test of the bee avoidance hypothesis.

Colour signals are the main floral trait for plant-pollinator communication. Owing to visual specificities, flower visitors exert different selective pressures on flower colour signals of plant communities. Although they evolved to attract pollinators, matching their visual sensitivity and colour preferences, floral signals may also evolve to avoid less efficient pollinators and antagonistic flower visitors. We evaluated evidence for the bee avoidance hypothesis in a Neotropical community pollinated mainly by bees and hummingbirds, the campo rupestre. We analysed flower reflectance spectra, compared colour variables of bee-pollinated flowers (bee-flowers; 244 species) and hummingbird-pollinated flowers (hummingbird-flowers; 39 species), and looked for evidence of bee sensorial exclusion in hummingbird-flowers. Flowers were equally contrasting for hummingbirds. Hummingbird-flowers were less conspicuous to bees, reflecting mainly long wavelengths and avoiding red-blind visitors. Bee-flowers reflected more short wavelengths, were more conspicuous to bees (higher contrasts and spectral purity) than hummingbird-flowers and displayed floral guides more frequently, favouring flower attractiveness, discrimination and handling by bees. Along with no phylogenetic signal, the differences in colour signal strategies between bee- and hummingbird-flowers are the first evidence of the bee avoidance hypothesis at a community level and reinforce the role of pollinators as a selective pressure driving flower colour diversity.

Does the Cave Environment Reduce Functional Diversity?

Caves are not colonised by all taxa present in the surface species pool, due to absence of light and the tendency to food limitation when compared to surface communities. Under strong species sorting during colonisation and later by the restrictive environmental filter, traits that are not adaptive in subterranean habitats may be filtered out. We tested whether cave communities were assembled by the restrictive regime propitiated by permanent darkness or by competitive exclusion due to resource scarcity. When compared to surface communities, the restrictive subterranean regime would lead to lower functional diversity and phenotypic clustering inside the caves, and the opposite should be expected in the case of competitive exclusion. Using isopods (Oniscidea) as model taxa, we measured several niche descriptors of taxa from surface and cave habitats, used a multivariate measure of functional diversity, and compared their widths. We found phenotypic overdispersion and higher functional diversity in cave taxa when compared to surface taxa. On the one hand, the dry climate outside of caves hampered the survival of several taxa and their ecological strategies, not viable under severe desiccation risk, culminating in the clustering of functional traits. In contrast, this restriction does not occur inside of caves, where isopods find favourable conditions under lower predation pressures and more amenable environmental parameters that allow occupation and subsequent diversification. Our results showed that, at least for some taxa, caves may not be such a harsh environment as previously thought. The high functional diversity we found inside caves adds an additional reason for the conservation of these sensitive environments.

Assessing community assembly along a successional gradient in the North Adriatic Karst with functional and phylogenetic distances.

Recently, two approaches to account for ecological differences in community composition have been developed: one based on trait differences (functional diversity) and another based on evolutionary history (phylogenetic diversity). Combining them allows an integrated view of processes structuring communities. The North Adriatic Karst is covered by species-rich grasslands, but land abandonment has resulted in replacement by woodlands. This creates a successional gradient along which environmental conditions may change, and different community assembly rules may apply. We sampled 56 plant communities in the Karst and used functional-phylogenetic distances to assess assembly along a successional gradient, from grasslands to shrublands and woodlands. We found 146 species, for which we measured functional traits and built a phylogenetic tree. The three successional stages were floristically different, with grasslands having species that are typical of harsher soil conditions and woodlands with species preferring milder soil conditions. All communities had higher functional than phylogenetic distances, implying that closely related species tended to be phenotypically dissimilar. When more importance was given to functional distances, most grasslands and some shrublands were underdispersed; when more importance was given to phylogenetic distances, only one grassland was underdispersed and one woodland was overdispersed. Combining functional and phylogenetic distances provided us with better estimates of ecological differences in a successional gradient, where environmental filters seem to be the dominant force in early stages and competitive exclusion becomes more important in later stages. Taking into account that sucessional stages are assembled by different rules is essential to predicting their behaviour under future environmental scenarios.

Are fire, soil fertility and toxicity, water availability, plant functional diversity, and litter decomposition related in a Neotropical savanna?

Understanding how biodiversity and ecosystem functioning respond to changes in the environment is fundamental to the maintenance of ecosystem function. In realistic scenarios, the biodiversity-ecosystem functioning path may account for only a small share of all factors determining ecosystem function. Here, we investigated the strength to which variations in environmental characteristics in a Neotropical savanna affected functional diversity and decomposition. We sought an integrative approach, testing a number of pairwise hypotheses about how the environment, biodiversity, and functioning were linked. We used structural equation modelling to connect fire frequency, soil fertility, exchangeable Al, water availability, functional diversity of woody plants, tree density, tree height, and litter decomposition rates in a causal chain. We found significant effects of soil nutrients, water availability, and Al on functional diversity and litter decomposition. Fire did not have a significant direct effect on functional diversity or litter decomposition. However, fire was connected to both variables through soil fertility. Functional diversity did not influence rates of litter decomposition. The mediated effects that emerged from pairwise interactions are encouraging not only for predicting the functional consequences of changes in environmental variables and biodiversity, but also to caution against predictions based on only environmental or only biodiversity change.

Protozooplankton and its relationship with environmental conditions in 13 water bodies of the Mogi-Guacu basin - SP, Brazil

Protozooplankton is an important component of the aquatic microbial food webs and its composition, density, and distribution reflect the chemical, physical, and biological aspects of the environment. Considering the scarce literature on freshwater protozoans in Brazil and on protozoan ecology in subtropical environments, we listed the ciliates and amoebae taxa found in 13 water bodies in São Paulo State and analyzed their abundance in relation to the environmental variables. We collected two samples in each environment, fixed immediately with mercuric chloride and stained with bromophenol blue. After microscopical analysis, 74 protozoan genera were identified and the Ciliophora were dominant in the majority of the environments. The Stichotrichia, represented mostly by the genus Halteria, occurred in all environments, and was the dominant subclass in five of them. The canonic correspondence analysis of the most frequent genera and the environmental variables showed that nitrite and nitrate were the variables that better explained the distribution of Limnostrombidium, Urotricha, and Vorticella. The densities of the genera Halteria, Coleps, and of the species Cinetochilum margaritaceum were positively affected by increasing concentrations of dissolved oxygen, particulate phosphate, conductivity, and temperature. C. margaritaceum were also negatively affected by increasing concentrations of nitrite and nitrate. Considering that we made only one sampling in each environment, the richness was high compared to the mean diversity of lakes in the São Paulo State. The Diogo Lake, located in an ecological reserve, was the richest one, confirming the need of more research on the biodiversity of more preserved environments.

O protozooplâncton é um componente importante da rede trófica microbiana de ambientes aquáticos e sua composição, densidade e distribuição refletem os aspectos físicos, químicos e bióticos do ambiente. Considerando a escassa literatura sobre protozoários de água doce no Brasil e sobre sua ecologia em ambientes subtropicais, inventariamos os táxons de ciliados e amebas em 13 corpos d'água do Estado de São Paulo e analisamos a variação na abundância dos gêneros/espécies de maior incidência em relação às variáveis ambientais. Coletamos duas amostras por ambiente, fixando-as com cloreto de mercúrio e corando-as com azul de bromofenol para posterior quantificação e identificação em microscópio ótico. Identificamos 74 gêneros de ciliados e amebas, e os Ciliophora dominaram na maioria dos ambientes. A subclasse Stichotrichia ocorreu em todos os ambientes, predominando em cinco deles, especialmente pela ocorrência o gênero Halteria. A Análise de Correspondência Canônica mostrou que as concentrações de nitrito e nitrato são as principais variáveis que explicam a distribuição dos gêneros Limnostrombidium, Urotricha e Vorticella. O aumento na concentração de oxigênio dissolvido, condutividade, temperatura e concentração de fosfato particulado afetou positivamente a densidade dos gêneros Halteria e Coleps e da espécie Cinetochilum margaritaceum, que foi ainda influenciada negativamente pelo aumento nas concentrações de nitrito e nitrato. Considerando-se que foi realizada apenas uma coleta, a riqueza de espécies foi alta quando comparada à média de taxa encontrada para corpos d'água do Estado de São Paulo. O ambiente mais rico, Lagoa do Diogo, localiza-se em uma estação ecológica, confirmando a necessidade de mais pesquisas sobre a diversidade em ambientes menos impactados.

The drivers of woody species richness and density in a Neotropical savannah.

Environmental filtering prevents species without certain attributes from occurring in local communities. Traits respond differently to different abiotic factors, assembling communities with varying composition along environmental gradients. Here, we measured proxies of soil fertility, disturbance by fire, response and physiological traits to assess how these variables interact to determine woody species richness and density in a Neotropical savannah. We explicitly incorporated our assumptions about how different abiotic filters influence different subsets of traits into a statistical model using structural equation modelling, yielding a more accurate representation of the assembly process. Fire had an effect on resistance traits, whereas soil fertility influenced physiological traits. Resistance traits explained both the richness and density of plots, whereas physiological traits explained only the density. Fewer fire events led to richer and denser plots. Similarly, areas with lower cation exchange capacity assembled less dense communities. Furthermore, we showed that structural equation modelling yielded a realistic representation of the bivariate interactions of distinct environmental filters with different subsets of traits.

The role of fire in structuring trait variability in Neotropical savannas.

Intraspecific trait variability plays a fundamental role in community structure and dynamics; however, few studies have evaluated its relative importance to the overall response of communities to environmental pressures. Since fire is considered a key factor in Neotropical savannas, we investigated to what extent the functional effects of fire in a Brazilian savanna occurs via intra- or interspecific trait variability. We sampled 12 traits in communities subjected to three fire regimes in the last 12 years: annual, biennial, and protected. To evaluate fire's relative effects, we fitted a general linear mixed models with species as random and fire as fixed factors, using: (1) all species in the communities (i.e., considering intra- and interspecific variabilities); (2) 18 species common to all fire regimes (i.e., intraspecific variability only); and (3) all species with their overall average trait values (i.e., interspecific variability only). We assessed the relative role of intra- or interspecific variability by comparing the significance of each trait in the three analyses. We also compared the within and between fire variabilities with a variance component analysis. Five traits presented larger intraspecific than interspecific variability, and the main effect of fire occurred at the intraspecific level. These results confirm that it is important to consider intraspecific variability to fully understand fire-prone communities. Moreover, trait variability was larger within than among fire regimes. Thus, fire may act more as an external filter, preventing some of the species from the regional pool from colonizing the cerrado, than as an internal factor structuring the already filtered cerrado communities.

O cerrado não é um bioma / The Brazilian cerrado is not a biome

É importante definirmos corretamente um termo, porque pode haver implicações práticas e imediatas. Dois conceitos importantes para serem definidos acuradamente são os de "cerrado" e de "bioma". Da maneira como vem sendo usado no Brasil, o conceito de bioma adquiriu erroneamente uma conotação florística. Na verdade, o conceito de bioma é similar ao de formação vegetal, mas leva em conta a associação da vegetação com a fauna e com os microrganismos. Assim, por um lado, o conceito de bioma é fisionômico e funcional, isto é, levam-se em conta a aparência geral da vegetação e aspectos como os ritmos de crescimento e reprodução; por outro, o conceito não é florístico, isto é, a afinidade taxonômica das espécies que aparecem em várias unidades de um mesmo bioma é irrelevante. Para sermos coerentes com toda a literatura internacional e usarmos o conceito de bioma acuradamente, devemos considerar o cerrado sensu lato como formado por três biomas: o campo tropical (campo limpo), a savana (campo sujo, campo cerrado e cerrado sensu stricto) e a floresta estacional (cerradão).

It is important to define correctly a given concept, because there may be immediate and practical implications. Two important concepts to be accurately defined are "cerrado" and "biome". As used in Brazil, the concept of biome has erroneously acquired a floristic meaning. As a matter of fact, the concept of biome is close to that of vegetation form, but takes into account the association between the vegetation and the animals and microorganisms. Thus, on the one hand, the concept of biome is physiognomic and functional, that is, it takes into account the general aspect of the vegetation, growth patterns, and reproduction patterns; on the other hand, it is not floristic, that is, the taxonomic affinities of the species that occur in different regions of the same biome are irrelevant. To be coherent with international literature and use accurately the concept of biome, we should consider the Brazilian cerrado as being composed of three biomes: tropical grassland (campo limpo), savanna (campo sujo, campo cerrado, and cerrado sensu stricto), and seasonal forest (cerradão).

Dispersal syndromes related to edge distance in cerrado sensu stricto fragments of central-western Brazil

The main selective forces affecting the fruiting strategies are related to the environment in which plants occur. As a savanna, microclimatic conditions should not vary in relation to distance from edge in cerrado sensu stricto fragments. Thus, we postulated that the importance of different dispersal syndromes would not vary towards the fragment core. Our aim was to test in four cerrado sensu stricto fragments in central Brazil whether the absolute density of anemo-, auto-, and zoochorous individuals varied in relation to edge distance. According to results, the absolute density of anemo-and autochorous individuals did not vary, whereas those of zoochorous individuals increased with edge distance, pointing out that there were other factors rather than abiotic conditions shifting zoochorous species to the interior of cerrado sensu stricto fragments.

As principais forças seletivas que afetam as estratégias de frutificação estão relacionadas ao ambiente em que as plantas ocorrem. Como fragmentos de cerrado sensu stricto são savânicos, as condições microclimáticas não devem variar em relação à distância da borda. Assim, postulamos que a importância das diferentes síndromes de dispersão não varia da borda ao interior de um fragmento de vegetação. O objetivo deste trabalho foi testar, em quatro fragmentos de cerrado sensu stricto (centro-oeste do Brasil), se a densidade absoluta de indivíduos anemo, auto e zoocóricos variava em função da distância da borda. Segundo nossos resultados, a densidade absoluta de indivíduos anemo e autocóricos não variou significativamente borda em direção ao interior dos fragmentos, enquanto que a dos indivíduos zoocóricos aumentou, indicando que existem outros fatores, que não as condições abióticas, deslocando as espécies zoocóricas para o interior dos fragmentos de cerrado sensu stricto.

Diversidades filogenética e funcional: novas abordagens para a Ecologia de comunidades / Phylogenetic and functional diversities: new approaches to community Ecology

Embora a diversidade pareça ser o conceito ecológico mais intuitivo, nenhuma definição consensual foi formulada. As medidas tradicionais de diversidade, que levam em conta apenas o número de espécies e suas contribuições relativas, têm se mostrado estimativas pouco preditivas da estrutura e do funcionamento das comunidades. Medidas de diversidade que incorporem informações sobre as relações filogenéticas das espécies ou suas características funcionais podem ser melhores do que as medidas tradicionais para muitas finalidades. Apresentamos uma pequena revisão das propriedades e aplicações de algumas medidas de diversidade. Enfatizamos aqui duas abordagens recentes e promissoras, as diversidades filogenética e funcional, que têm se mostrado mais sensíveis para detectar respostas das comunidades às mudanças ambientais. Na diversidade filogenética, as relações de parentesco entre as espécies são levadas em conta, enquanto que na diversidade funcional traços que devem ter relações com o funcionamento das comunidades são considerados. Discutimos ainda os desafios e as perspectivas para o uso dessas duas abordagens na ecologia.

Although diversity seems to be the most intuitive ecological concept, no consensual definition has been stated. Traditional diversity measures, which take into account only the number of species and their relative contribution, have little predictive power about the functioning of communities. Diversity measures that include information on phylogenetic relationships among species or their functional traits should be better than the traditional measures. We present a short review of the properties and applications of some diversity measures, emphasizing two recent and promising approaches, the phylogenetic and functional diversities, which have been shown to be more sensitive to detect responses of communities to environmental changes. In phylogenetic diversity, species relatedness is taken into account, whereas in functional diversity traits related with community functioning are considered. We also discuss challenges and perspectives for the use of these two approaches in ecology.

Vertical and horizontal distribution of pollination systems in cerrado fragments of central Brazil

In fragments of the cerrado, we determined the frequency of pollination systems and analyzed their spatial distribution. We placed 38 transects, sampling 2,280 individuals and 121 species. As expected in Neotropical regions, bee-pollination was the most frequent pollination system. We found a decrease in the frequency of plants pollinated by beetles towards the fragment interior. Similarly, we found significant variation in relation to height just for the bats; there was an increase in the frequency of plants pollinated by bats towards the higher heights. In general, we found no horizontal and vertical variation in the pollination systems, probably as consequence of the more open physiognomy of the cerrado vegetation.

As principais pressões seletivas nas estratégias de polinização originam principalmente do ambiente em que plantas ocorrem, como subdossel, dossel, borda ou interior de um fragmento. Diferentes condições ambientais aumentam as diferenças entre os nichos ecológicos e podem implicar diferenças nas proporções dos sistemas de polinização. Em fragmentos de cerrado, determinamos a freqüência dos sistemas de polinização e analisamos sua distribuição espacial. Lançamos 38 transecções aleatoriamente, amostrando 2.280 indivíduos e 121 espécies. Como esperado para regiões neotropicais, a polinização por abelhas foi o sistema de polinização mais freqüente. Encontramos uma diminuição na freqüência de plantas polinizadas por besouros em direção ao interior do fragmento. De modo similar, encontramos uma variação significativa em relação à altura somente para os morcegos, havendo um aumento na freqüência de plantas em direção a alturas mais altas. Em geral, não encontramos variações horizontais e verticais nos sistemas de polinização, provavelmente, como conseqüência da fisionomia mais aberta de cerrado.

The vascular flora of the cerrado in Emas National Park (Central Brazil): a savanna flora summarized

The Brazilian cerrado may be divided in two main phytogeographic sectors: one characterized by Piptocarpha rotundifolia, in which the Emas National Park (ENP) is located, and other characterized by Curatella americana. We carried out a floristic survey in ENP, which allowed an assessment of the taxonomic composition, taxa size, and similarity with other sites for the ENP's vascular flora. We compared the ENP's flora with southeastern outlying cerrado sites, also in the Piptocarpha sector, and with general floristic patterns in the cerrado vegetation. The distribution of species per family in ENP was significantly different from that obtained for each component of the general cerrado flora. The herbaceous component was characterized by an overproportion of Myrtaceae and an underproportion of Orchidaceae and Lythraceae; and the woody component, by an overproportion of Myrtaceae and Nyctaginaceae. When compared with outlying cerrado sites, the ENP was quite distinct, not only at species level, but also at family level.

O cerrado pode ser dividido em dois principais setores fitogeográficos: um caracterizado por Piptocarpha rotundifolia, em que o Parque Nacional das Emas (PNE) está localizado, e outro caracterizado por Curatella americana. Realizamos um levantamento florístico no PNE, que nos permitiu determinar a composição taxonômica de sua flora vascular e sua similaridade com outros sítios. Comparamos a flora do PNE com sítios disjuntos de cerrado, também no setor Piptocarpha, e com padrões florísticos gerais do cerrado. A distribuição de espécies por família foi significativamente diferente daquela obtida para cada componente da flora do cerrado. O componente herbáceo-subarbustivo se caracterizou pela super-representação de Myrtaceae e pela sub-representação de Orchidaceae e Lythraceae; e o componente arbustivo-arbóreo, pela super-representação de Myrtaceae e Nyctaginaceae. Quando comparado aos sítios disjuntos, o PNE se mostrou distinto, não só em nível específico, mas também em nível de famílias.